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The Role of the Medial Frontal Cortex in Cognitive Control
Abstract
Adaptive goal-directed behavior involves monitoring of ongoing actions and performance outcomes, and subsequent adjustments
of behavior and learning. We evaluate new findings in cognitive neuroscience concerning cortical interactions that subserve
the recruitment and implementation of such cognitive control. A review of primate and human studies, along with a meta-analysis
of the human functional neuroimaging literature, suggest that the detection of unfavorable outcomes, response errors, response
conflict, and decision uncertainty elicits largely overlapping clusters of activation foci in an extensive part of the posterior
medial frontal cortex (pMFC). A direct link is delineated between activity in this area and subsequent adjustments in performance.
Emerging evidence points to functional interactions between the pMFC and the lateral prefrontal cortex (LPFC), so that monitoring-related
pMFC activity serves as a signal that engages regulatory processes in the LPFC to implement performance adjustments.
... These studies identify the frontally localized error-related negativity (ERN) ERP component and fMRI activation in the dorsal anterior cingulate cortex (dACC) as error markers. However, other brain areas, such as the posterior cingulate cortex (PCC), are active during error monitoring and also express error-related fMRI activation (Ridderinkhof et al., 2004;Wittfoth et al., 2008). Several findings, mainly from studies with adults, show that the cingulate cortex is the most likely generator of the ERN activity (Agam et al., 2011;Herrmann et al., 2004;Tamnes et al., 2013). ...
... Neuroimaging studies using fMRI find strong activation in frontal brain areas on tasks requiring relatively high levels of conflict resolution. These areas include the dACC and other cingulo-opercular regions, dorsolateral prefrontal cortex (DLPFC), and other prefrontal regions (Carter et al., 1998;MacDonald et al., 2000;Ridderinkhof et al., 2004;Roe et al., 2021;Taylor et al., 2007;van Veen et al., 2001). The dACC is more active in various tasks requiring performance monitoring and error detection (Le et al., 2021;Nee et al., 2007;Ridderinkhof et al., 2004;Weiss & Luciana, 2022). ...
... These areas include the dACC and other cingulo-opercular regions, dorsolateral prefrontal cortex (DLPFC), and other prefrontal regions (Carter et al., 1998;MacDonald et al., 2000;Ridderinkhof et al., 2004;Roe et al., 2021;Taylor et al., 2007;van Veen et al., 2001). The dACC is more active in various tasks requiring performance monitoring and error detection (Le et al., 2021;Nee et al., 2007;Ridderinkhof et al., 2004;Weiss & Luciana, 2022). Performance monitoring and error detection mechanisms have been investigated also through event-related potential measures. ...
The ability to monitor performance during a goal-directed behavior differs among children and adults in ways that can be measured with several tasks and techniques. As well, recent work has shown that individual differences in error monitoring moderate temperamental risk for anxiety and that this moderation changes with age. We investigated age differences in neural responses linked to performance monitoring using a multimodal approach. The approach combined functional MRI and source localization of event-related potentials (ERPs) in 12-year-old, 15-year-old, and adult participants. Neural generators of two components related to performance and error monitoring, the N2 and ERN, lay within specific areas of fMRI clusters. Whereas correlates of the N2 component appeared similar across age groups, age-related differences manifested in the location of the generators of the ERN component. The dorsal anterior cingulate cortex (dACC) was the predominant source location for the 12-year-old group; this area manifested posteriorly for the 15-year-old and adult groups. A fMRI-based ROI analysis confirmed this pattern of activity. These results suggest that changes in the underlying neural mechanisms are related to developmental changes in performance monitoring.
... Дорсальная АСС и прилегающие к ней зоны mPFC активируются также при увеличении вероятности конфликта между несколькими вариантами возможных действий, например, в ситуации неопределенности или необходимости торможения непосредственных реакций. В обзоре [Ridderinkhof et al., 2004] на основе данных исследований ССП и фМРТ мозга человека, а также нейронной активности в мозге приматов выделены 4 типа процессов, с которыми связывают активность dACC. Это -детекция возможного конфликта будущей реакции с правильным ответом (pre-response conflict), возраста-ние неопределенности в исходе принятого решения (decision uncertainly), ошибочный ответ (response error) и получение экзогенной негативной обратной связи о совершенном действии. ...
... Связь dACC c процессом принятия решений, особенно в ситуации возможного конфликта при выборе стимула или реакции, определяет вовлечение этой области коры в процессы контроля когнитивной деятельности, о чем свидетельствуют многочисленные нейрокогнитивные исследования [см. обзоры Ridderinkhof et al., 2004;Heilbronner, Hayden, 2016], в том числе исследования с использованием так называемых «конфликтных» экспериментальных моделей. Согласно концепции М. Познера [Petersen, Posner, 2012] dACC является ключевой структурой одной из сетей внимания, а именно центральной управляющей сети. ...
В книге обобщены исследования коллектива авторов, посвященные актуальным проблемам мозгового обеспечения регуляции поведения и когнитивной деятельности в подростковом возрасте.
Морфологические, нейрофизиологические и нейропсихологические исследования, представленные в книге, свидетельствуют о существенных преобразованиях мозговой организации процессов регуляции познавательной деятельности и поведения у подростков. При этом направленность таких изменений — прогресс или снижение функциональных возможностей — различна для разных звеньев мозговых регуляторных систем и меняется с возрастом. Наиболее существенные изменения в подростковом возрасте претерпевают системы эмоционально-мотивационной регуляции, что в значительной степени увеличивает риск учебной и социальной дезадаптации.
Книга предназначена для физиологов, психологов, педагогов и студентов биологического, психолого-педагогического и клинико-психологического профиля.
... An ERP component capturing these processes may be the error-related negativity (ERN) [29][30][31][32][33] originating from medial frontal cortices. 34 The ERN reflects a comparison process between the desired and the achieved action outcome, 35 which is also required during continuous sensorimotor processing. 26 It is commonly accepted that such aspects of performance monitoring are necessary when adjusting action-to-action outcomes. ...
... 26 It is commonly accepted that such aspects of performance monitoring are necessary when adjusting action-to-action outcomes. 34 In addition, it is also mandatory to increase processing capacities to adapt complex movements when sensorimotor processes have to be adapted (because they deviate too much from the desired state). Such resource allocation processes are likely reflected by the P2 ERP component, [36][37][38] while adaptations in the mapping of the stimulus on the appropriate response are reflected by the P3 ERP component. ...
Monitoring actions is essential for goal-directed behaviour. However, as opposed to short-lasting, and regularly reinstating monitoring functions, the neural processes underlying continuous action monitoring are poorly understood. We investigate this using a pursuit-tracking paradigm. We show that beta band activity likely maintains the sensorimotor program, while theta and alpha bands probably support attentional sampling and information gating, respectively. Alpha and beta band activity are most relevant during the initial tracking period, when sensorimotor calibrations are most intense. Theta band shifts from parietal to frontal cortices throughout tracking, likely reflecting a shift in the functional relevance from attentional sampling to action monitoring. This study shows that resource allocation mechanisms in prefrontal areas and stimulus-response mapping processes in the parietal cortex are crucial for adapting sensorimotor processes. It fills a knowledge gap in understanding the neural processes underlying action monitoring and suggests new directions for examining sensorimotor integration in more naturalistic experiments.
... comparison to other prefrontal regions. For instance, while the IFG has been systematically linked to the retrieval and selection of information from memory Wagner, 2002, 2007;Bar, 2003;Thompson-Schill et al., 1997), the orbitofrontal gyrus may have instead a regulatory role of performance (Bar, 2003(Bar, , 2007Ridderinkhof et al., 2004a;. After activation of stimulus-specific information at perirhinal areas, it is thought that this region plays a role in adjusting and directing back predictions only on the objects relevant features to support the appropriate behaviour according to the task demands (Bar, 2003(Bar, , 2007. ...
... Moreover, there was no evidence of a typicality effect at IFG for older participants. Instead, the presumed reorganization of semantic connectivity (Dubossarsky et al., 2017;Wulff et al., 2018Wulff et al., , 2019 and/or the potential decline of neural responsivity in areas activated in younger adults (Bruffaerts et al., 2019;Hoffman and Morcom, 2018) seems to have elicited greater prefrontal control demands, particularly for more atypical concepts, possibly involving monitoring processes located at mOFG to redirect processing after adjusting predictions based on prior object information (Bar, 2003(Bar, , 2007Ridderinkhof et al., 2004a;. Interestingly, along with the top-down role of orbitofrontal areas, there is evidence of ATL as a crucial component in modulating semantic processing. ...
Effective use of conceptual knowledge engages semantic representation and control processes to access information in a goal-driven manner. Neuropsychological findings of patients presenting either degraded knowledge (e.g., semantic dementia) or disrupted control (e.g., semantic aphasia) converge with neuroimaging evidence from young adults, and delineate the neural segregation of representation and control mechanisms. However, there is still scarce research on the neurofunctional underpinnings of such mechanisms in healthy ageing. To address this, we conducted an fMRI study, wherein young and older adults performed a covert naming task of typical and atypical objects. Three main age-related differences were found. As shown by age group and typicality interactions, older adults exhibited overactivation during naming of atypical (e.g., avocado) relative to typical concepts in brain regions associated to semantic representation, including anterior and medial portions of left temporal lobe (respectively, ATL and MTG). This provides evidence for the reorganization of neural activity in these brain regions contingent to the enrichment of semantic repositories in older ages. The medial orbitofrontal gyrus was also overactivated, indicating that the processing of atypical concepts (relative to typical items) taxes additional control resources in the elderly. Increased activation in the inferior frontal gyrus (IFG) was observed in naming typical items (relative to atypical ones), but only for young adults. This suggests that naming typical items (e.g., strawberry) taxes more on control processes in younger ages, presumably due to the semantic competition set by other items that share multiple features with the target (e.g., raspberry, blackberry, cherry). Together, these results reveal the dynamic nature of semantic control interplaying with conceptual representations as people grow older, by indicating that distinct neural bases uphold semantic performance from young to older ages. These findings may be explained by neural compensation mechanisms coming into play to support neurocognitive changes in healthy ageing.
... Error-related brain activity has been studied using both electroencephalography (EEG) and functional magnetic resonance imaging (fMRI) approaches and is typically linked to the dorsal anterior cingulate cortex (dACC) and surrounding posterior medial frontal cortex (pMFC; Overbye et al., 2019) among other, more widely distributed regions (e.g., insula; Iannaccone et al., 2015). At the core of neural networks for error processing, the pMFC and its subregions have been implicated in error-related functions of performance monitoring, error-detection, and interference processing in youth and adults (Fitzgerald, Perkins, et al., 2010;Ridderinkhof et al., 2004). ...
... The association of greater error-related dACC activity with less OCS in a community sample of youth extends prior work demonstrating atypical pMFC response to errors in patients with OCD (Carrasco et al., 2013;Norman et al., 2019;Riesel et al., 2019). Consistent with prior literature, a main effect of errors was observed in the dACC and surrounding pMFC, including the SMA and pre-SMA (Norman et al., 2019;Ridderinkhof et al., 2004). Within the pMFC, the effect of OCS was localized to the dACC. ...
Background:
Subclinical obsessive-compulsive symptoms (OCS) are common in children, and increase risk for later onset of obsessive-compulsive disorder (OCD). In pediatric patients with OCD, neuroimaging research implicates altered neural mechanisms for error-processing, but whether abnormal brain response occurs with subclinical OCS remains poorly understood.
Methods:
Using functional magnetic resonance imaging (fMRI), 113 youth (8-18 years; 45 female) from a community sample were scanned during an error-eliciting Go/No-Go task. OCS were assessed dimensionally using the obsessive-compulsive subscale of the Child Behavior Checklist. The association between OCS scores and error-related brain activity was examined at the whole-brain level.
Results:
Lower OCS scores associated with stronger response to errors in dorsal anterior cingulate cortex (dACC), caudate, putamen, thalamus, and occipital cortex. Additionally, lower OCS related to higher capacity for inhibitory control, as indexed by greater accuracy on No-Go trials during fMRI scanning. The relationship between lower OCS and better accuracy on No-Go trials was mediated by greater error-related dACC activity.
Conclusions:
The inverse relationship between OCS and error-related activity in the dACC and extended cortical-striatal-thalamic circuitry may index an adaptive process by which subclinical OCS are minimized in youth. Further, these results identify an observable pattern of brain activity that tracks with subclinical OCS severity. Understanding the link between neural networks for error processing and the normal to abnormal range of OCS may pave the way for brain-based strategies to identify children who are more likely to develop OCD and enable the targeting of preventive strategies to reduce risk.
... The Stroop task examines ability to inhibit prepotent responses, an important aspect of executive function or cognitive control (Baddeley, 2012;Miyake et al., 2000;Ridderinkhof et al., 2004). In the typical version of this task, subjects have to name the color of the ink used in words, when the words are either congruent (i.e. the word RED in red ink) or incongruent (i.e. the word BLUE in red ink). ...
... Our finding of activation in a cluster encompassing the left dorsolateral, ventrolateral and rostral anterior cingulate cortex and supplementary motor area during Stroop task performance is in close agreement with a meta-analyses of activations during Stroop and related task performance in HC (Xu et al., 2016). This region has been linked to cognitive control (Gratton et al., 2018;Ridderinkhof et al., 2004), specifically its 'evaluative' component, responsible for monitoring response execution and signaling when adjustments are necessary, in contrast to its 'regulative' function, responsible for the generation of task goals and their maintenance during execution, involving the lateral prefrontal cortex and orbitofrontal cortex. ...
Introduction:
Few studies have examined the functional brain correlates of the performance of the Stroop task in bipolar disorder (BD). It is also not known whether it is associated with failure of de-activation in the default mode network, as has been found in studies using other tasks.
Methods:
Twenty-four BD patients and 48 age, sex and educationally estimated intellectual quotient (IQ) matched healthy subjects (HS) underwent a functional MRI during performance of the counting Stroop task. Task-related activations (incongruent versus congruent condition) and de-activations (incongruent versus fixation) were examined using whole-brain, voxel-based methodology.
Results:
Both the BD patients and the HS showed activation in a cluster encompassing the left dorsolateral and ventrolateral prefrontal cortex and the rostral anterior cingulate cortex and supplementary motor area, with no differences between them. The BD patients, however, showed significant failure of de-activation in the medial frontal cortex and the posterior cingulate cortex/precuneus.
Conclusions:
The failure to find activation differences between BD patients and controls suggests that the 'regulative' component of cognitive control remains intact in the disorder, at least outside episodes of illness. The failure of de-activation found adds to evidence documenting trait-like default mode network dysfunction in the disorder.
... The neurophysiology of stress sets the hypothalamus as a central component, in which the paraventricular nucleus is the main integrator of stressors, activating systems such as the sympathetic-adrenomedullar and hypothalamus-pituitary-adrenal axes (5). The brain structures actively involved in stress responses include the prefrontal cortex (6) and the amygdala, whose activity is also associated with emotional processing (7). Prefrontal projections to the amygdala (8), as well as hippocampus projections to the amygdala and prefrontal cortex (9) are involved as well. ...
... Note that EEG oscillations in the h band have been consistently reported as a sensitive correlate of emotion processing (94) and also in heart-tobrain communication (38). Our results show that preferential heart-to-brain communication occurs over the frontal and parietal cortical regions, consistently with a previous report on stress (6) and correlates of cognitive operations (95). ...
Dynamical information exchange between central and autonomic nervous systems, as referred to functional brain-heart interplay, occurs during emotional and physical arousal. It is well documented that physical and mental stress lead to sympathetic activation. Nevertheless, the role of autonomic inputs in nervous-system-wise communication under mental stress is yet unknown. In this study, we estimated the causal and bidirectional neural modulations between EEG oscillations and peripheral sympathetic and parasympathetic activities using a recently proposed computational framework for a functional brain-heart interplay assessment, namely the sympathovagal synthetic data generation model. Mental stress was elicited in 37 healthy volunteers by increasing their cognitive demands throughout three tasks associated with increased stress levels. Stress elicitation induced an increased variability in sympathovagal markers, as well as increased variability in the directional brain-heart interplay. The observed heart-to-brain interplay was primarily from sympathetic activity targeting a wide range of EEG oscillations, whereas variability in the efferent direction seemed mainly related to EEG oscillations in the gamma band. These findings extend current knowledge on stress physiology, which mainly referred to top-down neural dynamics. Our results suggest that mental stress may not cause an increase in sympathetic activity exclusively as it initiates a dynamic fluctuation within brain-body networks including bidirectional interactions at a brain-heart level. We conclude that directional brain-heart interplay measurements may provide suitable biomarkers for a quantitative stress assessment and bodily feedback may modulate the perceived stress caused by increased cognitive demand.
... Currently, there are two explanations for theta oscillatory activity. One suggests that theta oscillatory activity is associated with violations of expectations (Tzur and Berger 2009), as many studies have shown that increased theta oscillatory activity is associated with cognitive control (Botvinick et al. 2001), conflict detection (Ridderinkhof et al. 2004) and the calculations of prediction errors of action values (Matsumoto et al. 2007;Oliveira et al. 2007). Padrão et al. (2013) recorded electrophysiological responses to monetary gains and losses in extreme groups of anhedonic and nonanhedonic participants and found that participants with anhedonia showed a drastic increase in frontal medial theta power after receiving the maximum monetary gain. ...
Laboratory studies reveal that young women with premenstrual syndrome (PMS) often exhibit decreased reward processing during the late luteal phase. However, studies based on the self-reports find opposite results (e.g., higher craving for high-sweet-fat food). These differences may lie in the difference between the stimulus used and measuring the different aspects of the reward. The present study was designed to expand previous work by using a classic monetary reward paradigm, simultaneously examining the motivational (i.e., reward anticipation, “wanting”) and emotional (i.e., reward outcome, “liking”) components of reward processing in women with high premenstrual symptoms (High PMS). College female students in their early twenties with High PMS (n = 20) and low premenstrual symptoms (Low PMS, n = 20) completed a monetary incentive delay task during their late luteal phase when the premenstrual symptoms typically peak. Brain activities in the reward anticipation phase and outcome phase were recorded using the magnetoencephalographic (MEG) imaging technique. No group differences were found in various behavioral measurements. For the MEG results, in the anticipation phase, when High PMS participants were presented with cues that predicted the upcoming monetary gains, they showed higher event-related magnetic fields (ERFs) than when they were presented with neutral non-reward cues. This pattern was reversed in Low PMS participants, as they showed lower reward cue-elicited ERFs than non-reward cue-elicited ones (cluster mass = 2560, cluster size = 891, p = .03, corrected for multiple comparisons), mainly in the right medial orbitofrontal and lateral orbitofrontal cortex (cluster mass = 375, cluster size = 140, p = .03, corrected for multiple comparisons). More importantly, women with High PMS had an overall significantly higher level of ERFs than women with Low PMS (cluster mass = 8039, cluster size = 2937, p = .0009, corrected for multiple comparisons) in the bilateral precentral gyrus, right postcentral gyrus, and left superior temporal gyrus (right: cluster mass = 410, cluster size = 128, p = .03; left: cluster mass = 352, cluster size = 98, p = .05; corrected for multiple comparisons). In the outcome phase, women with High PMS showed significantly lower theta power than the Low PMS ones for the expected non-reward feedback in the bilateral temporal-parietal regions (cluster mass = 47620, cluster size = 18308, p = .01, corrected for multiple comparisons). These findings reveal that the severity of PMS might alter reward anticipation. Specifically, women with High PMS displayed increased brain activities to reward-predicting cues and increased action preparation after the cues appear.
... The medial frontal gyrus and orbital gyri both belong to the prefrontal cortex, which covers the anterior part of the frontal lobe and participates in various critical cognitive and emotional processing tasks. [43][44][45][46] Previous reports have shown that ALFF values significantly decreased in the medial frontal gyrus in several retinal diseases such as retinal detachment, 19 retinal vein occlusion patients, 23 and optic neuritis. 16 Our results of decreased medial frontal gyrus further confirmed these findings. ...
... The supplementary motor area is implicated in cognitive control, especially performance monitoring and behavioral planning (Bonini et al., 2014;Lemogne et al., 2011;Lim et al., 2015;Nachev et al., 2008). Maltreated individuals might have a more frequent need to evaluate and adjust performance to avoid mistakes and threat, having learned that mistakes could result in harsh punishment in abusive settings (Ridderinkhof et al., 2004). Hyperactivation of this key error-processing region in maltreated individuals may be abuse specific (Lim et al., 2015). ...
Gray matter (GM) abnormalities have been reported in both adults and children/adolescents with histories of childhood maltreatment (CM). As maturational and compensatory changes may occur over the life span, a comparison of effects in youth and adulthood may be informative regarding life-span effects of CM. Voxel-wise meta-analyses of whole-brain voxel-based morphometry (VBM) studies were conducted in all datasets and age-based subgroups respectively, followed by a quantitative meta-analytic comparison of the age-based subgroups. Thirty VBM studies (31 datasets) comprising 1264 individuals with CM (1004 adults and 260 children/adolescents) and 1201 non-maltreated controls (962 adults and 239 children/adolescents) were included. Maltreatment-exposed youth showed less GM in the cerebellum, and greater GM in bilateral middle cingulate/paracingulate gyri and bilateral visual cortex than maltreated adults. Opposite GM alterations in bilateral middle cingulate/paracingulate gyri were found in maltreatment-exposed adults (decreased) and children/adolescents (increased). Our findings demonstrate different patterns of GM changes in youth closer to maltreatment events than those seen later in life, suggesting detrimental effects of CM on the developmental trajectory of brain structure.
... A second highly relevant aspect is cognitive control. It allows to select contextually relevant information and organize/optimize information processing to reach goal-directed behavior (Ridderinkhof et al. 2004). However, individuals with SAUD are strongly impaired in inhibition (e.g., Quoilin et al. 2018). ...
Severe alcohol use disorder (SAUD) is among the most prevalent psychiatric disorders, leading to a wide range of deleterious consequences for the individual, their relatives and the society. Individuals suffering from SAUD are often tough to be more aggressive and violent. We reviewed the literature linking alcohol consumption and aggression to provide a synthesis of available evidence on this topic. In general, both acute alcohol consumption and SAUD are associated with an increase in the three subcomponents of aggression, namely anger, hostility and aggressive behaviors. We identified several mechanisms involved in the elicitation of aggression in SAUD, including hostility biases and the lack of emotional or cognitive control. Finally, we proposed treatments that specifically target these processes.
... While the ERN has been functionally linked to error/mismatch detection (Falkenstein et al., 2000), reinforcement learning (Holroyd & Coles, 2002), and/or postresponse conflict processing (Yeung et al., 2004), all theoretical models converge around the idea that the ERN reflects an early signal for the implementation of control mechanisms following action mistakes (Gehring et al., 2012). The ERN has been consistently linked to the activity of structures in the posterior medial frontal brain, including the anterior cingulate cortex and presupplementary motor area (Herrmann et al., 2004;Iannaccone et al., 2015;van Veen & Carter, 2002)-regions found to be implicated in situations requiring increased cognitive control, such as response conflict, performance errors, and decision uncertainty (Ridderinkhof et al., 2004;Ullsperger et al., 2014). Aside from theories linking the ERN to cognitive control mechanisms, other theoretical views have highlighted the relevance of motivational factors, positing that variations in ERN amplitude may reflect, at least in part, a motivational defensive response to signals of a potential threat (Hajcak & Foti, 2008;Weinberg et al., 2012). ...
The organization of the Hierarchical Taxonomy of Psychopathology (HiTOP) model provides unique opportunities to evaluate whether neural risk measures operate as indicators of broader latent liabilities (e.g., externalizing proneness) or narrower expressions (e.g., antisociality and alcohol abuse). Following this approach, the current study recruited a sample of 182 participants (54% female) who completed measures of externalizing psychopathology (also internalizing) and associated traits. Participants also completed three tasks (Flanker-No Threat, Flanker-Threat, and Go/No-Go tasks) with event-related potential (ERP) measurement. Three variants of two research domain criteria (RDoC)-based neurophysiological indicators-P3 and error-related negativity (ERN)-were extracted from these tasks and used to model two latent ERP factors. Scores on these two ERP factors independently predicted externalizing factor scores when accounting for their covariance with sex-suggesting distinct neural processes contributing to the broad externalizing factor. No predictive relation with the broad internalizing factor was found for either ERP factor. Analyses at the finer-grained level revealed no unique predictive relations of either ERP factor with any specific externalizing symptom variable when accounting for the broad externalizing factor, indicating that ERN and P3 index general liability for problems in this spectrum. Overall, this study provides new insights about neural processes in externalizing psychopathology at broader and narrower levels of the HiTOP hierarchy. (PsycInfo Database Record (c) 2023 APA, all rights reserved).
... We speculated that the dorsolateral prefrontal cortex (dlPFC) might be a key region involved in controlling Pavlovian bias. The dlPFC has been implicated in higher-level cognitive control and goal-directed actions [4,5,[10][11][12][13][14][15][16][17]. For example, dieters showed hyperactivation of the dlPFC when they successfully selected healthy food over tasty food [4]. ...
Previous literature suggests that a balance between Pavlovian and instrumental decision-making systems is critical for optimal decision-making. Pavlovian bias (i.e., approach toward reward-predictive stimuli and avoid punishment-predictive stimuli) often contrasts with the instrumental response. Although recent neuroimaging studies have identified brain regions that may be related to Pavlovian bias, including the dorsolateral prefrontal cortex (dlPFC), it is unclear whether a causal relationship exists. Therefore, we investigated whether upregulation of the dlPFC using transcranial current direct stimulation (tDCS) would reduce Pavlovian bias. In this double-blind study, participants were assigned to the anodal or the sham group; they received stimulation over the right dlPFC for 3 successive days. On the last day, participants performed a reinforcement learning task known as the orthogonalized go/no-go task; this was used to assess each participant's degree of Pavlovian bias in reward and punishment domains. We used computational modeling and hierarchical Bayesian analysis to estimate model parameters reflecting latent cognitive processes, including Pavlovian bias, go bias, and choice randomness. Several computational models were compared; the model with separate Pavlovian bias parameters for reward and punishment domains demonstrated the best model fit. When using a behavioral index of Pavlovian bias, the anodal group showed significantly lower Pavlovian bias in the punishment domain, but not in the reward domain, compared with the sham group. In addition, computational modeling showed that Pavlovian bias parameter in the punishment domain was lower in the anodal group than in the sham group, which is consistent with the behavioral findings. The anodal group also showed a lower go bias and choice randomness, compared with the sham group. These findings suggest that anodal tDCS may lead to behavioral suppression or change in Pavlovian bias in the punishment domain, which will help to improve comprehension of the causal neural mechanism.
... Xu et al. [97] found that the pre-supplementary motor area (pre-SMA) and the right inferior frontal gyrus (IFG) were jointly invoked by cognitive conflict and affective conflict. Accumulating evidence shows that the pre-SMA is involved in conflict control [39,51,66,75,76,90,91]. Both stimulation and impairment of the pre-SMA impact performance during conflict tasks [67,88,91]. ...
Studies have shown that there are overlapping neural bases for cognitive and affective conflict control, but whether the neural activity patterns caused by the two types of conflict are similar remains to be explored. The present study utilizes electroencephalogram (EEG) and functional magnetic resonance imaging (fMRI) to temporally and spatially analyze the differences between cognitive and affective conflict control. We employ a semantic conflict task which includes blocks of cognitive and affective judgements primed by conflicting and non-conflicting contexts. The results showed a typical neural conflict effect in the cognitive judgment blocks as reflected by greater amplitudes of P2, N400, and the late positive potential (LPP), as well as greater activation of the left pre-supplementary motor area (pre-SMA) and the right inferior frontal gyrus (IFG) in the conflict condition relative to the non-conflict condition. These patterns did not emerge in the affective judgments, but instead, showed reversed effects of the LPP and in the SMA. Taken together, these findings suggest that cognitive and affective conflict control result in different neural activity patterns. Full-text available at: https://www.sciencedirect.com/science/article/abs/pii/S0166432823002401
... Our finding that the dACC was involved in both emotional and domain-general conflict control accords with previous studies in which it was strongly engaged in non-emotional and emotional conflict control (Chechko et al., 2013;Godinez et al., 2016;Krug & Carter, 2010;Müller et al., 2011;Ochsner et al., 2009;Torres-Quesada et al., 2014). The dACC is an integrative function in behavioural control (Shackman et al., 2011) that detects conflict between a previous and new task in case of a task switch (Ridderinkhof et al., 2004) and produces ongoing behavioural modulation (Sheth et al., 2012) in the case of conflict adaptation (Botvinick et al., 2004). Accordingly, the dACC may provide a continuously updated prediction of ongoing cognitive demands while implementing behavioural adaptations to optimize performance. ...
The present study uses functional magnetic resonance image (fMRI) to examine
the overlapping and specific neural correlates of contextualized emotional conflict control and domain-general conflict control. During a performance on emotional and domain-general conflict tasks, conjunction analyses showed that neural areas distributed in the frontoparietal network were engaged in both processes, supporting the notion that similar neural mechanisms are implemented in these two types of control. Importantly, disjunction analyses revealed a broader neural recruitment of emotional conflict control compared to domain-general conflict control as shown by the possible lateralization of the lateral prefrontal cortex (lPFC), such that emotional conflict control significantly involved the left lPFC while domain-general conflict control seemly involved the right lPFC. Results of generalized psychophysiological interaction (gPPI) analyses further demonstrated that emotional conflict control, compared to domain-general conflict control, elicited broader synergistic activities in individuals' brain networks. Together, these findings offer novel and compelling neural evidence that furthers our understanding of the complex relationship between domain-general and emotional conflict control.
... An event-related potential (ERP) component investigated in the context of error processing is the error-related negativity (ERN), a negative-going frontocentral deflection that peaks around 100 ms after an erroneous response (Falkenstein et al., 1991;Falkenstein et al., 2000;Gehring et al., 1993Holroyd and Coles, 2002). The ERN appears to be generated in the mPFC, probably the ACC (Debener et al., 2005;Dehaene et al., 1994;Ridderinkhof et al., 2004;Taylor et al., 2007). ...
Errors elicit a negative, mediofrontal, event-related potential (ERP), for both own errors (error-related negativity; ERN) and observed errors (here referred to as observer mediofrontal negativity; oMN). It is unclear, however, if the action-monitoring system codes action valence as an all-or-nothing phenomenon or if the system differentiates between errors of different severity. We investigated this question by recording electroencephalography (EEG) data of pianists playing themselves (Experiment 1) or watching others playing (Experiment 2). Piano pieces designed to elicit large errors were used. While active participants’ ERN amplitudes differed between small and large errors, observers’ oMN amplitudes did not. The different pattern in the two groups of participants was confirmed in an exploratory analysis comparing ERN and oMN directly. We suspect that both prediction and action mismatches can be coded in action monitoring systems, depending on the task, and a need-to-adapt signal is sent whenever mismatches happen to indicate the magnitude of the needed adaptation.
... For example, researchers have consistently found that incongruent trials typically produce a larger fronto-central N2 than congruent stimuli in the Flanker task (Falkenstein et al., 1999;Gehring et al., 1992;Purmann et al., 2011). Moreover, findings from functional Magnetic Resonance Imaging (fMRI) studies and dipole source localization analyses show that the N2 is likely to reflect the activity of the anterior cingulate cortex (ACC) (Botvinick et al., 1999;Hinault et al., 2019;Nieuwenhuis et al., 2003), which has been found to be associated with cognitive control components such as conflict detection and response monitoring (Kopp et al., 1996;Ridderinkhof et al., 2004;van Veen & Carter, 2002;Yeung & Cohen, 2006), rather than novelty effects or motor inhibition per se (Huster et al., 2013). On the other hand, MFN is a negative deflection that occurs 0-100 ms after a behavioral response at frontal-central electrodes (Gehring & Knight, 2000;Johnson et al., 2008). ...
A number of psychological theories propose that deception involves more cognitive control than truth-telling. Over the last decades, event-related potentials (ERPs) have been used to unravel this question, but the findings are mixed. To address this controversy, two meta-analyses were conducted to quantify the results of existing studies reporting N2 or medial frontal negativity (MFN) associated with deception. In total, 32 papers consisting of 1091 participants were included, which yielded 32 effect sizes for N2 and 7 effect sizes for MFN. We found that deception was associated with a more negative N2 and MFN than truth-telling with medium and large effect sizes (r = .25 and .51, respectively). We also found that the deception paradigm modulated the results (p = .043), but found no evidence for publication bias. Our findings indicate that deception involves more cognitive control than truth-telling. Our review also identifies gaps in this literature, including a need for more ERP studies using spontaneous deception.
... The medial prefrontal cortex (mPFC) is associated with decision making (Euston et al., 2012), including detection errors (Holroyd et al., 2002) and cognitive control (Ridderinkhof et al., 2004), which is divided into anterior cingulate cortex (ACC), prelimbic cortex (PrL), and infralimbic cortex (IL), based on functional and anatomical characteristics (Heidbreder and Groenewegen, 2003;Sparta et al., 2014). As a cortical region, mPFC is made up of different neuronal types and projections, most of which are excitatory glutamatergic neurons, and a smaller proportion are inhibitory GABAergic neurons (Xu et al., 2019). ...
Hoarding disorder (HD) is a chronic disease that begins early in life and does not remission unless timely treated. A large number of factors affect the presentation of HD symptoms, including a strong possessive psychology of objects and neurocognitive functioning. However, the underlying neural mechanisms of the excessive hoarding behavior in HD are still unknown. Using viral infections and brain slice electrophysiology recordings, we found that increased glutamatergic neuronal activity and decreased GABAergic neuronal activity in medial prefrontal cortex (mPFC) accelerated the hoarding-like behavior in mice. Respectively, chemogenetic manipulation to reduce glutamatergic neuronal activity or enhance GABAergic neuronal activity could improve the hoarding-like behavioral response. These results reveal a critical role played by alterations in the activity of specific types of neurons in hoarding-like behavior, and that targeted therapies for HD may be possible by precisely modulating these types of neurons.
... Recent findings have repeatedly shown that event file coding processes are associated with inferior parietal structures (Kleimaker et al., 2020;Opitz et al., 2020;Takacs et al., 2020). The anterior cingulate cortex (ACC) also being associated with activity reflected by IC 20 is well-known to be associated with processes occurring in the Nogo-P3 time window (Albert et al., 2012;Bokura et al., 2001;Fallgatter et al., 2004;Gonzalez-Rosa et al., 2013;Huster et al., 2013), which fits to the general role of the ACC in behavioral monitoring processes (Ridderinkhof et al., 2004). Thus, the current neurophysiological findings derived through combining ICA, deep learning procedures, and source localization procedures provide evidence for concomitant processes being reflected in the Nogo-P3 time window. ...
Inhibitory control processes are an important aspect of executive functions and goal-directed behavior. However, the mostly correlative nature of neurophysiological studies was not able to provide insights which aspects of neural dynamics can best predict whether an individual is confronted with a situation requiring the inhibition of a response. This is particularly the case when considering the complex spatio-temporal nature of neural processes captured by EEG data. In the current study we ask whether independent spatial activity profiles in the EEG data are useful to predict whether an individual is confronted with a situation requiring response inhibition. We combine independent component analysis (ICA) with explainable artificial intelligence approaches (EEG-based deep learning) using data from a Go/Nogo task (N = 255 participants). We show that there are four dissociable spatial activity profiles important to classify Go and Nogo trials as revealed by deep learning. Of note, for all of these four independent activity profiles, neural activity in the time period between 300 and 550ms after stimulus presentation was most informative. Source localization analyses further revealed regions in the pre-central gyrus (BA6), the middle frontal gyrus (BA10), the inferior frontal gyrus (BA46) and the insular cortex (BA13) were associated with the isolated spatial activity profiles. The data suggest for concomitant processes being reflected in the identified time window. This has implications for the ongoing debate on the functional significance of event-related potential correlates of inhibitory control.
... It is important to highlight that these same areas of the social brain are also involved in mediating other aspects of cognition. The medial prefrontal cortex is involved in ToM functioning (Gallagher and Frith 2003) and decision-making (Ridderinkhof et al. 2004). The fusiform gyrus supports face processing (Kanwisher 2000). ...
Volumes of literature have been devoted to understanding how facial expressions of emotions are processed in autism. However, the nature of emotional processing outside the visual domain still remains unclear. Although emotional language incorporates a myri- ad of faculties, the current chapter highlights how individuals with autism process emotion words. Behavioral findings show that individuals are able to correctly identify basic emo- tion words. But neuroimaging data reveals that participants with autism fail to activate either motor or limbic systems in contrast to their typical counterparts. Furthermore, they also recruit language areas of both the hemispheres to a larger extent. These findings signi- fy differential integration of language, motor and emotion networks in autism even in the face of equivalent behavioral competence. Although the issue of emotion language process- ing requires further investigation, these studies indicate that at least high-functioning indi- viduals with autism can learn abstract emotional concepts in a rule-based manner that makes it difficult to decode emotions in the visual domain. Further, hypoactivity in the motor areas implies atypical motor embodiment of emotional concepts and aligns this do- main of research with documented motor deficits and anomalous emotional processing in the visual domain as part of the autistic symptomatology.
... These clusters connect with segregated networks (Figure 4b-d) that, however, overlap in the midcingulate cortex (Figure 4e). This region is considered an emerging target for neurostimulation (Downar et al., 2016;Mattavelli et al., 2022), because of its role in multiple facets of self-regulation and control, via performance monitoring, feedback detection and behavioural adjustments to improve performance based on both endogenous (e.g., goals) and exogenous (e.g., context) factors (Ridderinkhof et al., 2004). Importantly, this In particular, the adMCC is known for its crucial role in aspects of executive functioning targeted by cognitive training (Hoffstaedter et al., 2014;Margulies & Uddin, 2019;de la Vega et al., 2016), such as monitoring one's own and others' decision and outcomes (Apps et al., 2013;Rolls, 2019;Rushworth et al., 2004), error and novelty detection (Procyk et al., 2014), or task switching (Rushworth et al., 2002;Worringer et al., 2019). ...
While mounting evidence shows promising effects of brain training on cognitive functioning in healthy and pathological conditions, the spread of variable training approaches highlights the need to compare their efficacy and identify their neural correlates, representing possible targets for neuromodulation treatments. We performed coordinate-based functional magnetic resonance imaging meta-analyses to compare the neural correlates and the cognitive outcomes of cognitive (n = 22), physical (n = 22), and meditative (n = 20) training in healthy non-expert individuals. Pre/post-training cognitive metrics improved after cognitive and physical training, but their heterogeneity, or even the lack of these measurements in some studies, highlights the need of more structured protocols. Cognitive, physical, and meditative interventions increased brain activity in distinct fronto-medial areas likely mediating training effects on cognitive, action, and attentional control, respectively. The modular , training-specific, engagement of a region that is known to mediate feedback-based learning provides cues for boosting brain training via combined interventions that might jointly outperform their individual effects.
KEYWORDS: cognitive training, fMRI, medial prefrontal cortex, meditative training, meta-analysis, physical training, treatment
... However, we observed a signi cant difference in theta band power uctuations in the ON vs OFF condition, near the double support period in the left prefrontal cortex, right sensorimotor cortex, and right somatosensory cortex (Fig. 4). Studies have shown that power in medial and left prefrontal cortices may be associated with error detection [51,52] and adjustment [20,32,[53][54][55][56] during movement adaptation. In addition, the theta power increase has been associated with balance perturbations from primary sensory and motor cortices, anterior cingulate cortex, temporal cortex, and parietal cortex [22,23,57,58]. ...
Walking with an exoskeleton is a complex task that requires human and machine integration. Previous studies focused on metabolic consumption, muscle activations, kinetic and kinematic changes, and adaption during exoskeleton-assisted walking. However, limited information is available on cortical changes and adaptations during walking with an exoskeleton. Our study aims to better understand human cortical responses and adaptation to walking with an ankle exoskeleton. We included healthy, novice users without prior exoskeleton experience and collected EEG (electroencephalography), EMG (electromyography), and full body motion capture while walking at a speed of 1.2m/s. When experiencing exoskeleton-assisted walking for the first time, subjects showed reduced joint range of motion and stride length along with increased step width, even while muscle activation was decreased. The cortical response also follows the decreased balance, showing theta power increase in the left prefrontal, right sensorimotor, and right somatosensory cortex. Over time, the results showed increased balance, greater reduction in muscle activity, and reduced cortical engagement while walking with exoskeleton assistance. Yet, we also observed adaptation to walking with exoskeleton frames over time. These findings could provide information to develop more effective and intuitive augmentation devices that are better able to integrate with human movements.
... ERN and PEP come from different sources; the N1 potential of PEP is localized in the supplementary motor area (SMA) [11,[49][50][51], while the ERN appears in the cingulate cortex (especially ACC) [29,[52][53][54][55][56]. Although both areas are localized in the pMFC, the research indicated that pMF acts as a 'neural alarm' and is responsive to a broad range of elicitors such as performance monitoring and adjustments, the detection of an error, anomalies, discrepancies, adverse outcomes, and decision uncertainty [30,[57][58][59]. ...
Objective:
The maintenance of balance is a complicated process in the human brain, which involves multisensory processing such as somatosensory and visual processing, motor planning and execution. It was shown that a specific cortical activity called perturbation-evoked potential (PEP) appears in the electroencephalogram (EEG) during balance perturbation. PEPs are primarily recognized by the N1 component with a negative peak localized in frontal and central regions. There has been a doubt in balance perturbation studies whether the N1 potential of perturbation is elicited due to error processing in the brain. The objective of this study is to test whether the brain perceives postural instability as a cognitive error by imposing two types of perturbations consisting of erroneous and correct perturbations.
Approach:
We conducted novel research to incorporate the experiment designs of both error and balance studies. To this end, participants encountered errors during balance perturbations at rare moments in the experiment. We induced errors by imposing perturbations to participants in the wrong directions and an erroneous perturbation was considered as a situation when the participant was exposed to an opposite direction of the expected/informed one. In correct perturbations, participants were tilted to the same direction, as they were informed. We analyzed the two conditions in time, time-frequency, and source domains.
Main results:
We showed that two error-related neural markers were derived from the EEG responses, including error positivity (Pe), and error-related alpha suppression (ERAS) during erroneous perturbations. Consequently, early neural correlates of perturbation cannot be interpreted as error-related responses. We discovered distinct patterns of conscious error processing; both Pe and ERAS are associated with conscious sensations of error.
Significance:
Our findings indicated that early cortical responses of balance perturbation are not associated with neural error processing of the brain, and errors induce distinct cortical responses that are distinguishable from brain dynamics of N1 potential.

.
... There is an ongoing debate as to the processes the ERN signal indexes: error detection or prediction of error-likelihood, 44-46 detection of response conflict, 47,48 action monitoring, [49][50][51][52] or increase in cognitive control 53 (see review in the study by Shalgi & Deouell 54 ). Studies analyzing single-unit recordings identify both dorsal anterior cingulate cortex (dACC) and pre-supplementary motor area (pre-SMA) as contributors to the ERN 9,49 (see also the study by Sajad et al. 55 measuring supplementary eye field in macaques), with pre-SMA neurons responding earlier than dACC neurons. ...
Some decisions make a difference, but most are arbitrary and inconsequential, like which of several identical new pairs of socks should I wear? Healthy people swiftly make such decisions even with no rational reasons to rely on. In fact, arbitrary decisions have been suggested as demonstrating "free will". However, several clinical populations and some healthy individuals have significant difficulties in making such arbitrary decisions. Here, we investigate the mechanisms involved in arbitrary picking decisions. We show that these decisions, arguably based on a whim, are subject to similar control mechanisms as reasoned decisions. Specifically, error-related negativity (ERN) brain response is elicited in the EEG following change of intention, without an external definition of error, and motor activity in the non-responding hand resembles actual errors both by its muscle EMG temporal dynamics and by the lateralized readiness potential (LRP) pattern. This provides new directions in understanding decision-making and its deficits.
... Evaluation and comparison of performance outcomes with internal standards are critical to optimizing future behaviors [3,43]. Outcome evaluation facilitates reinforcement learning and is involved in performance monitoring [21,45]. ...
It has been suggested that maladaptive perfectionists are more prone to concern over their performance outcomes than adaptive perfectionists. Performance outcome evaluation is reflected in the amplitude of feedback-related negativity (FRN) in brain electroencephalography (EEG). Hence, the amplitude of the FRN after receiving unfavorable feedback indicating a negative performance outcome may reflect personality characteristics. In other words, EEG could be a better marker of personality characteristics than self-report measures. However, the FRN component has not yet been investigated between different types of perfectionists. In the present study, group differences in the FRN were examined between two groups of adaptive and maladaptive perfectionists and a group of non-perfectionists during a monetary gambling task. We observed a larger FRN amplitude for adaptive perfectionists than for maladaptive perfectionists. This finding is consistent with previous reports that reward prediction error is reflected in the amplitude of the FRN. This difference in FRN could be interpreted as the pessimistic outcome expectation biases in maladaptive perfectionists.
... This specific activation of the ACC in ambiguous social settings as indicated in the present results aligns with previous findings. These findings attribute an integrative weighting function accountable for controlling adaptive social cognition mechanisms as well as mediating the value of social stimuli to the ACC [24][25][26][27][28][29]66,69,70]. This includes a personal cost-benefit evaluation and an estimate of the motivation of others [67,71]. ...
This study represents a follow-up event-related potential (ERP) analysis of a prior investigation. The previous results showed that participants had most negative-tending ERPs in the mid-frontal brain region during exposure to neutral emotion pictures (compared to negative and positive pictures) while being accompanied by a significant other person (social presence condition). The present analysis aimed at investigating potential sex differences related to this phenomenon. Female and male participants’ brain activity data from the previous study were analyzed separately for one representative mid-frontal electrode location selected on the basis of having the highest significance level. As a result, only female participants showed significantly more negative-tending potentials in response to neutral pictures, compared to both other emotion categories (positive and negative) in the social presence condition. This was not found in male participants. The respective ERP effect was most dominant at 838 ms post stimulus onset, which is slightly later than the effect found in the prior study. However, this result is interpreted as evidence that the general effect from the prior study can be understood as a largely female phenomenon. In line with the prior study, the present results are interpreted as a predominantly female activation in the mid-frontal brain region in response to neutral picture stimuli while being accompanied by a significant other person (social presence condition). Although only speculative, this would align with previous studies demonstrating sex-related hormonal and structural differences in the anterior cingulate cortex (ACC). In general, ACC activation has been associated with an integrative weighting function in ambiguous social settings, which makes sense given the ambiguous nature of neutral pictures in combination with a social presence condition.
... This network is activated during controlled processing of complex information, such as prediction and evaluation of multiple outcomes with high relational complexity (Kroger et al., 2002;Vincent et al, 2008). The dorsomedial SFG is also involved in complex cognitive processing, including performance monitoring, action selection and switching in preparation of responses (Ridderinkhof et al., 2004;Rushworth et al., 2004). ...
Exposure to altered g-levels causes unusual sensorimotor demands that must be dealt with by the brain. This study aimed to investigate whether fighter pilots, who are exposed to frequent g-level transitions and high g-levels, show differential functional characteristics compared to matched controls, indicative of neuroplasticity. We acquired resting-state functional magnetic resonance imaging data to assess brain functional connectivity (FC) changes with increasing flight experience in pilots and to assess differences in FC between pilots and controls. We performed whole-brain exploratory and region-of-interest (ROI) analyses, with the right parietal operculum 2 (OP2) and the right angular gyrus (AG) as ROIs. Our results show positive correlations with flight experience in the left inferior and right middle frontal gyri, and in the right temporal pole. Negative correlations were observed in primary sensorimotor regions. We found decreased whole-brain functional connectivity of the left inferior frontal gyrus in fighter pilots compared to controls and this cluster showed decreased functional connectivity with the medial superior frontal gyrus. Functional connectivity increased between the right parietal operculum 2 and the left visual cortex, and between the right and left angular gyrus in pilots compared to controls. These findings suggest altered motor, vestibular, and multisensory processing in the brains of fighter pilots, possibly reflecting coping strategies to altered sensorimotor demands during flight. Altered functional connectivity in frontal areas may reflect adaptive cognitive strategies to cope with challenging conditions during flight. These findings provide novel insights into brain functional characteristics of fighter pilots, which may be of interest to humans traveling to space.
... are necessary to achieve one's action goals (Holroyd & Yeung, 2012;Ridderinkhof, Ullsperger, Crone, & Nieuwenhuis, 2004). Also, the N e seems to be associated with affective-motivational aspects of error processing. ...
The literature on narcissism suggests two contradictory ways how highly narcissistic individuals deal with their failures: They might avoid consciously recognising their failures to protect their ego or they might vigilantly turn towards their failures to process cues that are important for maintaining their grandiosity. We tried to dissolve these contradictory positions by studying event-related potential components of error processing and their variations with narcissism. With a speeded go/no-go task, we examined how the error-related negativity (Ne; reflecting an early, automatic processing stage) and the error positivity (Pe; associated with conscious error detection) vary with Admiration and Rivalry, two narcissism dimensions, under ego-threatening conditions. Using multilevel models, we showed that participants with high Rivalry displayed higher Ne amplitudes suggesting a heightened trait of defensive reactivity. We did not find variations of either narcissism dimension with the Pe, which would have pointed to weaker error awareness. Thus, our results only supported the second position: a heightened vigilance to errors in narcissism at early, rather automatic processing stages.
Cognitive models state social anxiety (SA) involves biased cognitive processing that impacts what is learned and remembered within social situations, leading to the maintenance of SA. Neuroscience work links SA to enhanced error monitoring, reflected in error-related neural responses arising from mediofrontal cortex (MFC). Yet, the role of error monitoring in SA remains unclear, as it is unknown whether error monitoring can drive changes in memory, biasing what is learned or remembered about social situations. Thus, we developed a novel paradigm to investigate the role of error-related MFC theta oscillations (associated with error monitoring) and memory biases in SA. EEG was collected while participants completed a novel Face-Flanker task, involving presentation of task-unrelated, trial-unique faces behind target/flanker arrows on each trial. A subsequent incidental memory assessment evaluated memory biases for error events. Severity of SA symptoms were associated with greater error-related theta synchrony over MFC, as well as between MFC and sensory cortex. SA was positively associated with memory biases for error events. Consistent with a mechanistic role in biased cognitive processing, greater error-related MFC-sensory theta synchrony during the Face-Flanker predicted subsequent memory biases for error events. Our findings suggest high SA individuals exhibit memory biases for error events, and that this behavioral phenomenon may be driven by error-related MFC-sensory theta synchrony associated with error monitoring. Moreover, results demonstrate the potential of a novel paradigm to elucidate mechanisms underlying relations between error monitoring and SA.
The term, Mental Optometry , is newly developed concept that can be used to describe the interplay between mind, brain, and sensory interpretations. Taken from the premise of behavioral optometry and research explaining body orientation to physical field of vision, what we see or perceive with our mind’s eye, emotions and behaviors will also follow in the same manner. While not explicitly referred to in such a manner, cognitive, cognitive behavioral, and cognitive bias formation theories imply such a concept as being foundational to their systems. Mental Optometry arms the theorist and practitioner with a neurobiological empowered understanding of mood, emotion, thought, and interpretations of visual stimuli such that therapeutic interventions can be developed to assist patients in recognizing and altering skewed interpretations of what they think they see (the mind’s eye) – imagery that may deleteriously support negative cognitions leading to negative mood states.
Background:
Cocaine use disorder is associated with cognitive deficits that reflect dysfunctional processing across neural systems. As there are currently no approved medications, treatment centres around behavioural interventions which have only short-term efficacy. This suggests that behavioural interventions are not sufficient by themselves to maintain abstinence in cocaine use disorder patients. Self-control, which includes the regulation of attention, is critical for dealing with many daily challenges that would benefit from medication interventions that can ameliorate cognitive neural disturbances.
Methods:
To address this important clinical gap, we conducted a randomized, double-blind, placebo controlled, crossover design study in patients with cocaine use disorder (n = 23) and healthy control participants (n = 28). We assessed the modulatory effects of acute atomoxetine (40 mg) on attention and conflict monitoring, and their associated neural activation and connectivity correlates during performance on the Eriksen Flanker task. The Eriksen Flanker task examines basic attentional processing using congruent stimuli, and the effects of conflict monitoring and response inhibition using incongruent stimuli; the latter of which necessitates the executive control of attention.
Results:
We found that atomoxetine improved task accuracy only in the cocaine group, but modulated connectivity within distinct brain networks in both groups during congruent trials. During incongruent trials, the cocaine group showed increased task-related activation in the right inferior frontal and anterior cingulate gyri, as well as greater network connectivity compared with the control group, across treatments.
Conclusions:
This supports a modulatory effect of acute atomoxetine on attention and associated connectivity in cocaine use disorder.
Trial name:
Noradrenergic modulation of regulatory control and the function of related brain systems in cocaine dependence (Investigating the effects of Atomoxetine on Brain and Behaviour).
Registration number:
13999.
Background:
Goal-directed decision-making is a central component of the broader reward and motivation system, and requires the ability to dynamically integrate both positive and negative feedback from the environment in order to maximize rewards and minimize losses over time. Altered decision-making processes, in which individuals fail to consider the negative consequences of their decisions on both themselves and others, may play a role in driving antisocial behaviour.
Aim:
The main study aim was to investigate possible differences in loss and risk aversion across matched patients, all with a schizophrenia spectrum disorder (SSD), but who varied according to whether they had a history of serious interpersonal violence or not, and a sample of healthy controls with no history of violence.
Results:
The sample included 14 forensic and 21 non-forensic patients with SSD, and 41 healthy controls. Among the three decision-making variables under investigation, risk aversion was the only significant predictor of membership of the three groups, with greater risk aversion among non-forensic patients with SSD compared to healthy controls. No differences were observed across groups in loss aversion and choice consistency.
Conclusions:
This evidence suggests a new potential treatment target for rehabilitative measures aimed at achieving functional improvements in patients with SSD by selectively leveraging the neuro-cognitive processing of reward.
There is accumulating evidence for positive effects of green spaces on mental and brain health. Here we investigated whether differentiating the types of green spaces may be relevant. On longitudinal data of children ( N = 95) from the Netherlands, we quantified the link between green space exposure at home from birth onwards and MRI brain structure at 12.5 years. We differentiated between green space resulting from trees versus open green spaces and also associated visibility of sky (sky view factor) with brain structure (200 m buffer around home address). We observed a positive association between grey matter volume in different prefrontal clusters and green open space coverage as well as sky view, but a negative association within prefrontal clusters for tree cover density. Most importantly, in the medial prefrontal cortex, the only region in which all three analyses overlapped, the visibility of sky was the most important predictor. Our findings advance knowledge on health-promoting, evidence-based urban planning.
Background:
Cognitive impairment is the most common clinical manifestation of ischemic leukoaraiosis (ILA), but the underlying neurobiological pathways have not been well elucidated. Recently, it was thought that ILA is a "disconnection syndrome". Disorganized brain connectome were considered the key neuropathology underlying cognitive deficits in ILA patients.
Objective:
We aimed to detect the disruption of network hubs in ILA patients using a new analytical method called voxel-based eigenvector centrality (EC) mapping.
Methods:
Subjects with moderate to severe white matters hyperintensities (Fazekas score ≥3) and healthy controls (HCs) (Fazekas score = 0) were included in the study. The resting-state functional magnetic resonance imaging and the EC mapping approach were performed to explore the alteration of whole-brain network connectivity in ILA patients.
Results:
Relative to the HCs, the ILA patients exhibited poorer cognitive performance in episodic memory, information processing speed, and executive function (all ps < 0.0125). Additionally, compared with HCs, the ILA patients had lower functional connectivity (i.e., EC values) in the medial parts of default-mode network (i.e., bilateral posterior cingulate gyrus and ventral medial prefrontal cortex [vMPFC]). Intriguingly, the functional connectivity strength at the right vMPFC was positively correlated with executive function deficit in the ILA patients.
Conclusion:
The findings suggested disorganization of the hierarchy of the default-mode regions within the whole-brain network in patients with ILA and advanced our understanding of the neurobiological mechanism underlying executive function deficit in ILA.
The syndrome of apathy has generated increasing interest in recent years as systematic evaluations have revealed its high prevalence and strong negative impact on quality of life across a wide range of neurological and psychiatric conditions. However, although several theoretical models have been proposed to account for various aspects of the condition, understanding of this syndrome is still incomplete. One influential model has proposed that apathy might be described as a quantitative reduction of goal-directed behaviour in comparison to an individual's prior level of functioning. Persistence of activity defined as the capacity to continue with a task - sometimes in the face of setbacks, high levels of difficulty or fatigue - is a crucial but understudied aspect of goal-directed behaviour. Surprisingly, it has not been investigated yet in the context of apathy. Here, we provide an overview of theoretical and experimental aspects of persistence in effort that might assist to develop methods for the investigation of persistence in human behaviour, particularly within the pathologic context of apathy.
Background
Mild cognitive impairment (MCI) depicts a transitory phase between healthy elderly and the onset of Alzheimer's disease (AD) with worsening cognitive impairment. Some functional MRI (fMRI) research indicated that the frontoparietal network (FPN) could be an essential part of the pathophysiological mechanism of MCI. However, damaged FPN regions were not consistently reported, especially their interactions with other brain networks. We assessed the fMRI-specific anomalies of the FPN in MCI by analyzing brain regions with functional alterations.
Methods
PubMed, Embase, and Web of Science were searched to screen neuroimaging studies exploring brain function alterations in the FPN in MCI using fMRI-related indexes, including the amplitude of low-frequency fluctuation, regional homogeneity, and functional connectivity. We integrated distinctive coordinates by activating likelihood estimation, visualizing abnormal functional regions, and concluding functional alterations of the FPN.
Results
We selected 29 studies and found specific changes in some brain regions of the FPN. These included the bilateral dorsolateral prefrontal cortex, insula, precuneus cortex, anterior cingulate cortex, inferior parietal lobule, middle temporal gyrus, superior frontal gyrus, and parahippocampal gyrus. Any abnormal alterations in these regions depicted interactions between the FPN and other networks.
Conclusion
The study demonstrates specific fMRI neuroimaging alterations in brain regions of the FPN in MCI patients. This could provide a new perspective on identifying early-stage patients with targeted treatment programs.
Systematic review registration
https://www.crd.york.ac.uk/prospero/display_record.php?ID=CRD42023432042 , identifier: CRD42023432042.
Unawareness of memory deficits is an early manifestation in patients with Alzheimer's disease (AD), which often delays diagnosis. This intriguing behavior constitutes a form of anosognosia, whose neural mechanisms remain largely unknown. We hypothesized that anosognosia may depend on a critical synaptic failure in the error-monitoring system, which would prevent AD patients from being aware of their own memory impairment. To investigate, we measured event-related potentials (ERPs) evoked by erroneous responses during a word memory recognition task in two groups of amyloid positive individuals with only subjective memory complaints at study entry: those who progressed to AD within the five-year study period (PROG group), and those who remained cognitively normal (CTRL group). A significant reduction in the amplitude of the positivity error (Pe), an ERP related to error awareness, was observed in the PROG group at the time of AD diagnosis (vs study entry) in intra-group analysis, as well as when compared with the CTRL group in inter-group analysis, based on the last EEG acquisition for all subjects. Importantly, at the time of AD diagnosis, the PROG group exhibited clinical signs of anosognosia, overestimating their cognitive abilities, as evidenced by the discrepancy scores obtained from caregiver/informant vs participant reports on the cognitive subscale of the Healthy Aging Brain Care Monitor. To our knowledge, this is the first study to reveal the emergence of a failure in the error-monitoring system during a word memory recognition task at the early stages of AD. This finding, along with the decline of awareness for cognitive impairment observed in the PROG group, strongly suggests that a synaptic dysfunction in the error-monitoring system may be the critical neural mechanism at the origin of unawareness of deficits in AD.
Substantial evidence indicates that feedback processing not only varies with the valence of feedback, but is also highly dependent on contextual factors. Even so, the influence of prior outcome history on current outcome evaluation is far from clear. To investigate this issue, we conducted two event-related potential (ERP) experiments using a modified gambling task whereby each trial was associated with two consequences. In experiment 1, two instances of feedback indicated participant performance on two dimensions of a single decision, within a trial. In experiment 2, participants made two decisions in each trial, and then received two instances of feedback. We examined the feedback-related negativity (FRN) as an index of feedback processing. When both instances of feedback were relevant to the same trial (intra-trial), the FRN to the second was affected by the valence of the immediately previous feedback: The FRN was amplified to losses following wins. This was observed in both experiment 1 and experiment 2. When two instances of feedback were relevant to two different trials (inter-trial), the effect of immediately previous feedback on the FRN was inconsistent. In experiment 1 there was no effect of feedback from the previous trial on the FRN. However, in Experiment 2 there was an effect of inter-trial feedback on the FRN that was opposite to the effect of intra-trial feedback: The FRN was amplified when losses followed losses. Taken together, the findings suggest that the neural systems involved in reward processing dynamically and continuously integrate preceding feedback for the evaluation of present feedback.
Sensory responses of cortical neurons are more discriminable when evoked on a baseline of desynchronized spontaneous activity, but cortical desynchronization has not generally been associated with more accurate perceptual decisions. Here we show that mice perform more accurate auditory judgements when activity in the auditory cortex is elevated and desynchronized before stimulus onset, but only if the previous trial was an error, and that this relationship is occluded if previous outcome is ignored. We confirmed that the outcome-dependent effect of brain state on performance is neither due to idiosyncratic associations between the slow components of either signal, nor to the existence of specific cortical states evident only after errors. Instead, errors appear to gate the effect of cortical state fluctuations on discrimination accuracy. Neither facial movements nor pupil size during the baseline were associated with accuracy, but they were predictive of measures of responsivity, such as the probability of not responding to the stimulus or of responding prematurely. These results suggest that the functional role of cortical state on behavior is dynamic and constantly regulated by performance monitoring systems.
The Cambridge Handbook of Computational Cognitive Sciences is a comprehensive reference for this rapidly developing and highly interdisciplinary field. Written with both newcomers and experts in mind, it provides an accessible introduction of paradigms, methodologies, approaches, and models, with ample detail and illustrated by examples. It should appeal to researchers and students working within the computational cognitive sciences, as well as those working in adjacent fields including philosophy, psychology, linguistics, anthropology, education, neuroscience, artificial intelligence, computer science, and more.
Despite the many mistakes we make while speaking, people can effectively communicate because we monitor our speech errors. However, the cognitive abilities and brain structures that support speech error monitoring are unclear. There may be different abilities and brain regions that support monitoring phonological speech errors versus monitoring semantic speech errors. We investigated speech, language, and cognitive control abilities that relate to detecting phonological and semantic speech errors in 41 individuals with aphasia who underwent detailed cognitive testing. Then, we used support vector regression lesion symptom mapping to identify brain regions supporting detection of phonological versus semantic errors in a group of 76 individuals with aphasia. The results revealed that motor speech deficits as well as lesions to the ventral motor cortex were related to reduced detection of phonological errors relative to semantic errors. Detection of semantic errors selectively related to auditory word comprehension deficits. Across all error types, poor cognitive control related to reduced detection. We conclude that monitoring of phonological and semantic errors relies on distinct cognitive abilities and brain regions. Furthermore, we identified cognitive control as a shared cognitive basis for monitoring all types of speech errors. These findings refine and expand our understanding of the neurocognitive basis of speech error monitoring.
Purpose of Review
Impulsivity is a core feature and a risk factor of Internet Gaming Disorder (IGD). Hence, studies started examining the neural mechanisms that underlie this impulsivity. However, the big picture of such mechanisms is not clear. This paper seeks to survey recent cognitive neuroscience research on IGD and impulsivity and provide a synthesized view.
Recent Findings
Research has indicated that individuals with IGD have a greater degree of impulsivity than healthy controls (HC) and recreational Internet gaming users (RGU). This increased impulsivity has been associated with dysfunction or structural changes in the frontal lobe, striatum, amygdala, and insula, as well as the functional connectivity attributes between these areas. Nevertheless, there are some conflicting conclusions that should be explored further.
Summary
Studies have revealed that impulsivity is a risk factor for IGD, and that impulsivity facets are linked to IGD both behaviorally and neurobiologically. Additionally, the functional connections between the frontal lobe, striatum, amygdala, and insula could underlie the link between high impulsivity and IGD.
Intermittent fasting (IF) is an ecological strategy to control various metabolic disorder symptoms, but its protective effect on type 1 diabetes (T1D)-induced cognitive dysfunction and the underlying mechanisms remain poorly defined. Herein, we examined the efficacy of IF in altering the behaviors and brain metabolome in T1D mice and investigated the potential molecular mechanisms. We demonstrated that IF remarkably improved frontal cortical-dependent memory in T1D mice and reduced the loss of neuronal cells. Metabolomics and targeted mass spectrometry assay showed that IF reprogrammed the frontal cortical metabolome composition, including activated the aspartate and glutamate pathway and reversed glycerophospholipid and sphingolipid depositions in T1D mice. Mechanistically, IF attenuated the levels of oxidative stress proteins, such as NOX2, NOX4, 8-OHdG, 4-HNE, and inhibited the levels of pro-apoptotic factors Bax and cleaved Caspase-3, finally improved the memory ability of T1D mice. In vitro studies confirmed the protective effect of the supplemented N-acetylaspartate, a pivotal metabolite involved in IF-regulated T1D-induced cognitive dysfunction, in high glucose-stimulated SH-SY5Y cells by eliminating toxic lipids accumulation, oxidative stress and apoptosis. To conclude, the frontal cortical metabolites mediated the protective effects of IF against T1D-induced cognitive dysfunction by attenuating oxidative stress and apoptotic signaling. Thus, IF can be a potential therapeutic strategy for T1D-induced cognitive dysfunction.
The prefrontal cortex is included in a neuronal system that includes the basal ganglia, the thalamus, and the cerebellum. Most of the higher and more complex motor, cognitive, and emotional behavioral functions are thought to be found primarily in the frontal lobes. Insufficient connectivity between the medial prefrontal cortex (mPFC) and other regions of the brain that are distant from each other involved in top-down information pro- cessing rely on the global integration of data from multiple input sources and enhance low-level perception processes (bottom-up information processing). The reduced deactivation in mPFC and in the rest of the Default Network during global task processing is consistent with the integrative modulatory role served by the mPFC. We stress the importance of understanding the degree to which sensory and movement anomalies in individuals with autism spectrum disorder (ASD) can contribute to social impairment. Further investigation on the neurobiological basis of sensory symptoms and its relationship to other clinical features found in ASD is required Treatment perhaps should not be first behaviorally-based but rather based on facilitating sensory motor development
Prenatal opioid exposure is one consequence of the opioid epidemic, but effects on child development remain poorly understood. There is emerging evidence that children exposed to opioids in utero exhibit elevated emotional and behavioral problems, which may be partially due to alterations in cognitive control. Using multiple methods (i.e., neuropsychological, behavioral, and event-related potential [ERP] assessments), the present study examined differences in emotional, behavioral, and cognitive control difficulties in preschool-aged children with (n = 21) and without (n = 23) prenatal opioid exposure (Mage = 4.30, SD = 0.77 years). Child emotional and behavioral problems were measured with a caregiver questionnaire, indicators of cognitive control were measured using developmentally appropriate behavioral (i.e., delay discounting, Go/No-Go) and neuropsychological (i.e., Statue) tasks, and electroencephalogram was recorded to error and correct responses in a Go/No-Go task. ERP analyses focused on the error-related negativity (ERN), an ERP that reflects error monitoring, and correct-response negativity (CRN), a component reflecting performance monitoring more generally. Opioid exposure was associated with elevated difficulties across domains and a blunted ERN, reflecting altered cognitive control at the neural level, but groups did not significantly differ on behavioral measures of cognitive control. These result replicate prior studies indicating an association between prenatal opioid exposure and behavioral problems in preschool-aged children. Further, our findings suggest these differences may be partially due to children with prenatal opioid exposure exhibiting difficulties with cognitive control at the neural level. The ERN is a potential target for future research and intervention efforts to address the sequelae of prenatal opioid exposure.
Error-related electroencephalographic potentials have been used for decades to develop theoretical models of response monitoring processes, study altered cognitive functioning in clinical populations, and more recently, to improve the performance of brain-computer interfaces. However, the vast majority of this research relies on discrete behavioral responses that confound error detection, response cancellation, error correction, and post-error cognitive and affective processes. By contrast, the present study demonstrates a novel, complementary method for isolating the functional correlates of error-related electroencephalographic responses using single-trial kinematic analyses of cursor trajectories and a stepwise time-locking analysis. The results reveal that the latency of the ERN, Pe, and medial-frontal theta oscillations are all strongly positively correlated with the latency at which an initiated error response is canceled, as indicated by the peak deceleration of the initiated movement prior to a corrective response. Results are discussed with respect to current theoretical models of error-related brain potentials and potential relevance to clinical applications.
Human society encompasses diverse social influences, and people experience events differently and may behave differently under such influence, including in forming an impression of others. However, little is known about the underlying neural relevance of individual differences in following others' opinions or social norms. In the present study, we designed a series of tasks centered on social influence to investigate the underlying relevance between an individual's degree of social conformity and their neural variability. We found that individual differences under the social influence are associated with the amount of inter-trial electroencephalogram (EEG) variability over multiple stages in a conformity task (making face judgments and receiving social influence). This association was robust in the alpha band over the frontal and occipital electrodes for negative social influence. We also found that inter-trial EEG variability is a very stable, participant-driven internal state measurement and could be interpreted as mindset instability. Overall, these findings support the hypothesis that higher inter-trial EEG variability may be related to higher mindset instability, which makes participants more vulnerable to exposed external social influence. The present study provides a novel approach that considers the stability of one's endogenous neural signal during tasks and links it to human social behaviors.
Inattention to one’s on-going task leads to well-documented cognitive, behavioral, and physiological consequences. At the same time, the reliable association between mind-wandering and negative mood has suggested that there are affective consequences to task inattention as well. We examined this potential relationship between inattention and mood in the following study. Six hundred and fifty-five participants completed self-report questionnaires related to inattentive thinking (i.e., attentional lapses, daydreaming, mindfulness, rumination, reflection, worry, postevent processing, inattentiveness, and counterfactual thinking), a questionnaire about depressive symptoms, and a questionnaire about anxiety symptoms. First, an exploratory factor analysis was conducted to identify potential underlying constructs of types of inattentive thinking. Using ordinary least squares extraction and Oblimin rotation, a three-factor model demonstrated suitable fit, broadly representing mind-wandering/inattentive consequences, repetitive negative thinking, and reflective/introspective thinking. Second, after eliminating measures that did not strongly load on any factor, structural equation modeling was conducted and found that the relationship between mind-wandering and depression was partially explained by repetitive negative thinking, whereas the relationship between mind-wandering and anxiety was fully explained by repetitive negative thinking. The present findings suggest that understanding how inattentive thoughts are interrelated not only influences mood and affect but also reveals important considerations of intentionality, executive functioning, and qualitative styles of these thoughts.
Maintenance of a relatively high vigilance level is deemed to be important yet challenging when performing monotonous tasks with rare critical events that require crucial decision-making to avert catastrophic failures. With the rise in automation levels, we expect this problem to grow. In this chapter, the dynamics of sustained attention during naturalistic monotonous tasks will be presented. Various factors related to the nature of the task, stimuli, and individuals that contribute to the changes in vigilance are explored in detail. We also discuss methodological approaches for the assessment of vigilance decrement using neurological and physiological signals. Recent advances in real-time neuroimaging and neurofeedback have also encouraged researchers to investigate various strategies for vigilance enhancement in naturalistic tasks. In this light, several experimental studies that were designed to study vigilance decrement and enhancement during naturalistic tasks are examined. Key directions for future work in vigilance enhancement research are also proposed.
Summary Goal-directed behaviour depends on keeping relevant information in mind (working memory) and irrelevant information out of mind (behavioural inhibition or interference resolution). Prefrontal cortex is essential for working memory and for interference resolution, but it is unknown whether these two mental abilities are mediated by common or distinct prefrontal regions. To address this question, functional MRI was used to identify brain regions activated by separate manipulations of working memory load and interference within a single task (the Sternberg item recognition paradigm). Both load and interference manipulations were associated with performance decrements. Subjects were unaware of the interference manipulation. There was a high degree of overlap between the regions activated by load and interference, which included bilateral ventrolateral and dorsolateral prefrontal cortex, anterior insula, anterior cingulate and parietal cortex. Critically, no region was activated exclusively by interference. Several regions within this common network exhibited a brain–behaviour
A database of positron-emission-tomography studies published between January 1993 and November 1996 was created to address
several questions regarding the function and connectivity of the human anterior cingulate cortex (ACC). Using this database,
we have previously reported on the relationship between behavioural variables and the probability of blood-flow response in
distinct subdivisions of the ACC. The goal of the current analysis was to discover which areas of the frontal cortex show
increased blood-flow co-occurring consistently with increased blood-flow in the ACC. Analyses of the frequency distributions
of peaks in the ACC and the remaining frontal cortex (FC) yielded several important findings. First, FC peaks in the precentral
gyrus, superior frontal gyrus, middle frontal gyrus, inferior frontal gyrus, medial frontal gyrus and orbitomedial frontal
gyri were more frequent in subtractions that also yielded a peak in the ACC than in those that did not yield an ACC peak.
Second, regional differences in the frequency distribution of these FC peaks were observed when the ACC peaks were subdivided
into the rostral versus caudal ACC and supracallosal versus subcallosal ACC. Peaks in the precentral gyrus and in the vicinity
of the supplementary motor area were more prevalent in subtractions with co-occurring peaks in the caudal than with the rostral
ACC. Peaks in the middle frontal gyrus were more frequent in subtractions with co-occurring peaks in the paralimbic part of
the supracallosal ACC, relative to the subcallosal or limbic supracallosal ACC. These observations are consistent with known
differences in the anatomic connectivity in these cortical regions, as defined in non-human primates. Further analyses of
the influence of behavioural variables on the relationships between the ACC and other regions of the frontal cortex suggested
that this type of meta-analysis may provide testable hypotheses about functional and effective connectivity within the human
frontal lobe.
Most natural actions are chosen voluntarily from many possible choices. An action is often chosen based on the reward that it is expected to produce. What kind of cellular activity in which area of the cerebral cortex is involved in selecting an action according to the expected reward value? Results of an analysis in monkeys of cellular activity during the performance of reward-based motor selection and the effects of chemical inactivation are presented. We suggest that cells in the rostral cingulate motor area, one of the higher order motor areas in the cortex, play a part in processing the reward information for motor selection.
Intelligent behaviour requires self-control based on the consequences of actions. The countermanding task is designed to study self-control; it requires subjects to withhold planned movements in response to an imperative stop signal, which they can do with varying success. In humans, the medial frontal cortex has been implicated in the supervisory control of action. In monkeys, the supplementary eye field in the dorsomedial frontal cortex is involved in producing eye movements, but its precise function has not been clarified. To investigate the role of the supplementary eye field in the control of eye movements, we recorded neural activity in macaque monkeys trained to perform an eye movement countermanding task. Distinct groups of neurons were active after errors, after successful withholding of a partially prepared movement, or in association with reinforcement. These three forms of activation could not be explained by sensory or motor factors. Our results lead us to put forward the hypothesis that the supplementary eye field contributes to monitoring the context and consequences of eye movements.
The orbitofrontal cortex (OFC) is implicated in emotion and emotion-related learning. Using event-related functional magnetic resonance imaging (fMRI), we measured brain activation in human subjects doing an emotion-related visual reversal-learning task in which choice of the correct stimulus led to a probabilistically determined 'monetary' reward and choice of the incorrect stimulus led to a monetary loss. Distinct areas of the OFC were activated by monetary rewards and punishments. Moreover, in these areas, we found a correlation between the magnitude of the brain activation and the magnitude of the rewards and punishments received. These findings indicate that one emotional involvement of the human orbitofrontal cortex is its representation of the magnitudes of abstract rewards and punishments, such as receiving or losing money.
The prefrontal cortex has long been suspected to play an important role in cognitive control, in the ability to orchestrate thought and action in accordance with internal goals. Its neural basis, however, has remained a mystery. Here, we propose that cognitive control stems from the active maintenance of patterns of activity in the prefrontal cortex that represent goals and the means to achieve them. They provide bias signals to other brain structures whose net effect is to guide the flow of activity along neural pathways that establish the proper mappings between inputs, internal states, and outputs needed to perform a given task. We review neurophysiological, neurobiological, neuroimaging, and computational studies that support this theory and discuss its implications as well as further issues to be addressed
Dorsal anterior cingulate cortex (dACC) is a brain region that subserves cognition and motor control, but the mechanisms of these functions remain unknown. Human neuroimaging and monkey electrophysiology studies have provided valuable insights, but it has been difficult to link the two literatures. Based on monkey single-unit recordings, we hypothesized that human dACC is comprised of a mixture of functionally distinct cells that variously anticipate and detect targets, indicate novelty, influence motor responses, encode reward values, and signal errors. As an initial test of this conceptualization, the current event-related functional MRI study used a reward-based decision-making task to isolate responses from a subpopulation of dACC cells sensitive to reward reduction. As predicted, seven of eight subjects showed significant (P < 10(-4)) dACC activation when contrasting reduced reward (REDrew) trials to fixation (FIX). Confirmatory group analyses then corroborated the predicted ordinal relationships of functional MRI activation expected during each trial type (REDrew > SWITCH > CONrew > or = FIX). The data support a role for dACC in reward-based decision making, and by linking the human and monkey literatures, provide initial support for the existence of heterogeneity within dACC. These findings should be of interest to those studying reward, cognition, emotion, motivation, and motor control.
Event-related functional magnetic resonance imaging was used to measure blood oxygenation level-dependent responses in 13 young healthy human volunteers during performance of a probabilistic reversal-learning task. The task allowed the separate investigation of the relearning of stimulus-reward associations and the reception of negative feedback. Significant signal change in the right ventrolateral prefrontal cortex was demonstrated on trials when subjects stopped responding to the previously relevant stimulus and shifted responding to the newly relevant stimulus. Significant signal change in the region of the ventral striatum was also observed on such reversal errors, from a region of interest analysis. The ventrolateral prefrontal cortex and ventral striatum were not significantly activated by the other, preceding reversal errors, or when subjects received negative feedback for correct responses. Moreover, the response on the final reversal error, before shifting, was not modulated by the number of preceding reversal errors, indicating that error-related activity does not simply accumulate in this network. The signal change in this ventral frontostriatal circuit is therefore associated with reversal learning and is uncontaminated by negative feedback. Overall, these data concur with findings in rodents and nonhuman primates of reversal-learning deficits after damage to ventral frontostriatal circuitry, and also support recent clinical findings using this task.
The authors present a unified account of 2 neural systems concerned with the development and expression of adaptive behaviors: a mesencephalic dopamine system for reinforcement learning and a "generic" error-processing system associated with the anterior cingulate cortex. The existence of the error-processing system has been inferred from the error-related negativity (ERN), a component of the event-related brain potential elicited when human participants commit errors in reaction-time tasks. The authors propose that the ERN is generated when a negative reinforcement learning signal is conveyed to the anterior cingulate cortex via the mesencephalic dopamine system and that this signal is used by the anterior cingulate cortex to modify performance on the task at hand. They provide support for this proposal using both computational modeling and psychophysiological experimentation.
The anterior cingulate cortex (ACC) is a critical component of the human mediofrontal neural circuit that monitors ongoing
processing in the cognitive system for signs of erroneous outcomes. Here, we show that the consumption of alcohol in moderate
doses induces a significant deterioration of the ability to detect the activation of erroneous responses as reflected in the
amplitude of brain electrical activity associated with the ACC. This impairment was accompanied by failures to instigate performance
adjustments after these errors. These findings offer insights into how the effects of alcohol on mediofrontal brain function
may result in compromised performance.
The present study employed event-related fMRI and EEG to investigate the biological basis of the cognitive control of behavior. Using a GO/NOGO task optimized to produce response inhibitions, frequent commission errors, and the opportunity for subsequent behavioral correction, we identified distinct cortical areas associated with each of these specific executive processes. Two cortical systems, one involving right prefrontal and parietal areas and the second regions of the cingulate, underlay inhibitory control. The involvement of these two systems was predicated upon the difficulty or urgency of the inhibition and each was employed to different extents by high- and low-absent-minded subjects. Errors were associated with medial activation incorporating the anterior cingulate and pre-SMA while behavioral alteration subsequent to errors was associated with both the anterior cingulate and the left prefrontal cortex. Furthermore, the EEG data demonstrated that successful response inhibition depended upon the timely activation of cortical areas as predicted by race models of response selection. The results highlight how higher cognitive functions responsible for behavioral control can result from the dynamic interplay of distinct cortical systems.
Human anterior cingulate function has been explained primarily within a cognitive framework. We used functional MRI experiments with simultaneous electrocardiography to examine regional brain activity associated with autonomic cardiovascular control during performance of cognitive and motor tasks. Using indices of heart rate variability, and high- and low-frequency power in the cardiac rhythm, we observed activity in the dorsal anterior cingulate cortex (ACC) related to sympathetic modulation of heart rate that was dissociable from cognitive and motor-related activity. The findings predict that during effortful cognitive and motor behaviour the dorsal ACC supports the generation of associated autonomic states of cardiovascular arousal. We subsequently tested this prediction by studying three patients with focal damage involving the ACC while they performed effortful cognitive and motor tests. Each showed abnormalities in autonomic cardiovascular responses with blunted autonomic arousal to mental stress when compared with 147 normal subjects tested in identical fashion. Thus, converging neuroimaging and clinical findings suggest that ACC function mediates context-driven modulation of bodily arousal states.
The precise role of orbitofrontal cortex (OFC) in affective processing is still debated. One view suggests OFC represents stimulus reward value and supports learning and relearning of stimulus-reward associations. An alternate view implicates OFC in behavioral control after rewarding or punishing feedback. To discriminate between these possibilities, we used event-related functional magnetic resonance imaging in subjects performing a reversal task in which, on each trial, selection of the correct stimulus led to a 70% probability of receiving a monetary reward and a 30% probability of obtaining a monetary punishment. The incorrect stimulus had the reverse contingency. In one condition (choice), subjects had to choose which stimulus to select and switch their response to the other stimulus once contingencies had changed. In another condition (imperative), subjects had simply to track the currently rewarded stimulus. In some regions of OFC and medial prefrontal cortex, activity was related to valence of outcome, whereas in adjacent areas activity was associated with behavioral choice, signaling maintenance of the current response strategy on a subsequent trial. Caudolateral OFC-anterior insula was activated by punishing feedback preceding a switch in stimulus in both the choice and imperative conditions, indicating a possible role for this region in signaling a change in reward contingencies. These results suggest functional heterogeneity within the OFC, with a role for this region in representing stimulus-reward values, signaling changes in reinforcement contingencies and in behavioral control.
Consensus is emerging that the medial frontal lobe of the brain is involved in monitoring performance, but precisely what is monitored remains unclear. A saccade-countermanding task affords an experimental dissociation of neural signals of error, reinforcement, and conflict. Single-unit activity was monitored in the anterior cingulate cortex of monkeys performing this task. Neurons that signaled errors were found, half of which responded to the omission of earned reinforcement. A further diversity of neurons signaled earned or unexpected reinforcement. No neurons signaled the form of conflict engendered by interruption of saccade preparation produced in this task. These results are consistent with the hypothesis that the anterior cingulate cortex monitors the consequences of actions.
Conflict monitoring by the anterior cingulate cortex (ACC) has been posited to signal a need for greater cognitive control,
producing neural and behavioral adjustments. However, the very occurrence of behavioral adjustments after conflict has been
questioned, along with suggestions that there is no direct evidence of ACC conflict-related activity predicting subsequent
neural or behavioral adjustments in control. Using the Stroop color-naming task and controlling for repetition effects, we
demonstrate that ACC conflict-related activity predicts both greater prefrontal cortex activity and adjustments in behavior,
supporting a role of ACC conflict monitoring in the engagement of cognitive control.
In our event-related functional magnetic resonance imaging (fMRI) experiment, participants learned to select between two response options by trial-and-error, using feedback stimuli that indicated monetary gains and losses. The results of the experiment indicate that error responses and error feedback activate the same region of dorsal anterior cingulate cortex, suggesting that this region is sensitive to both internal and external sources of error information.
A neglected question regarding cognitive control is how control processes might detect situations calling for their involvement. The authors propose here that the demand for control may be evaluated in part by monitoring for conflicts in information processing. This hypothesis is supported by data concerning the anterior cingulate cortex, a brain area involved in cognitive control, which also appears to respond to the occurrence of conflict. The present article reports two computational modeling studies, serving to articulate the conflict monitoring hypothesis and examine its implications. The first study tests the sufficiency of the hypothesis to account for brain activation data, applying a measure of conflict to existing models of tasks shown to engage the anterior cingulate. The second study implements a feedback loop connecting conflict monitoring to cognitive control, using this to simulate a number of important behavioral phenomena.
Humans can monitor actions and compensate for errors. Analysis of the human event-related brain potentials (ERPs) accompanying errors provides evidence for a neural process whose activity is specifically associated with monitoring and compensating for erroneous behavior. This error-related activity is enhanced when subjects strive for accurate performance but is diminished when response speed is emphasized at the expense of accuracy. The activity is also related to attempts to compensate for the erroneous behavior.
Some years ago we described a negative (Ne) and a later positive (Pe) deflection in the event-related brain potentials (ERPs) of incorrect choice reactions [Falkenstein, M., Hohnsbein, J., Hoormann, J., Blanke, L., 1990. In: Brunia, C.H.M., Gaillard, A.W.K., Kok, A. (Eds.), Psychophysiological Brain Research. Tilburg Univesity Press, Tilburg, pp. 192–195. Falkenstein, M., Hohnsbein, J., Hoormann, J., 1991. Electroencephalography and Clinical Neurophysiology, 78, 447–455]. Originally we assumed the Ne to represent a correlate of error detection in the sense of a mismatch signal when representations of the actual response and the required response are compared. This hypothesis was supported by the results of a variety of experiments from our own laboratory and that of Coles [Gehring, W.J., Goss, B., Coles, M.G.H., Meyer, D.E., Donchin, E., 1993. Psychological Science 4, 385–390. Bernstein, P.S., Scheffers, M.K., Coles, M.G.H., 1995. Journal of Experimental Psychology: Human Perception and Performance 21, 1312–1322. Scheffers, M.K., Coles, M.G.H., Bernstein, P., Gehring, W.J., Donchin, E., 1996. Psychophysiology 33, 42–54]. However, new data from our laboratory and that of Vidal et al. [Vidal, F., Hasbroucq, T., Bonnet, M., 1999. Biological Psychology, 2000] revealed a small negativity similar to the Ne also after correct responses. Since the above mentioned comparison process is also required after correct responses it is conceivable that the Ne reflects this comparison process itself rather than its outcome. As to the Pe, our results suggest that this is a further error-specific component, which is independent of the Ne, and hence associated with a later aspect of error processing or post-error processing. Our new results with different age groups argue against the hypotheses that the Pe reflects conscious error processing or the post-error adjustment of response strategies. Further research is necessary to specify the functional significance of the Pe.
Two monkeys were trained to lift a lever with wrist extension in response to a visual cue. The animals were chronically implanted with electrodes for recording transcortical field potentials. Unique 'error' potentials were observed in the anterior cingulate cortical area 24 during a transitional learning-stage when the animals were uncertain about fulfilling the required task appropriately. At that stage (III) about 40-60% of the movements were made appropriately within the required cue-time. Error potentials followed inappropriate lever lifts made with wrist movements that were self-paced and hence non-rewarded, but such potentials did not follow appropriately made, i.e. visually initiated, and rewarded lifts.
Several areas on the medial surface of the frontal lobe in both monkeys and humans, including the supplementary motor area and specific areas within the ventral bank of the cingulate sulcus called the cingulate motor areas, have been implicated in the initiation and execution of skiled movements. These areas project directly to the motor cortex and spinal cord, and, on this basis alone, can be considered premotor areas. The present study investigated whether these premotor areas are specific targets of prefrontal cortical projections in the rhesus monkey and thereby provide links between this association cortex and motor effector pathways. Circumscribed injections of wheat germ agglutinin-conjugated horseradish peroxidase were placed into different cytoarchitectonic subdivisions of prefrontal cortex, and resultant retrograde and anterograde labeling examined with respect to designated premotor targets. Conversely, injections were also made in the supplementary and cingulate motor areas and labeled cells and terminals charted in the prefrontal cortex.
A principal finding in this study is the identification of multiple prefrontal regions that project to the supplementary motor area, the cingulate motor areas, or both. Areas 46, 8a, 9, 11, and 12 are reciprocally connected with an area of the superior frontal gyrus in or near the supplementary motor area at its rostral margin. A smaller constellation of prefrontal areas, areas 46, 8a, and 11, is reciprocally connected with portions of cingulate cortex that have been classified as premotor arm and/or leg representations (Hutchins et al., Exp Brain Res 71:667–672, 1988). In accordance with numerous previous reports, prefrontal areas 46, 8a, 9, 10, 11, and 12 are reciprocally connected with “nonmotor” subdivisions of cingulate cortex.
The results presented here specify the corticocortical connections by which prefrontal cortex may influence motor output.
Our goal in this review is to provide an anatomical framework for the analysis of the motor functions of the medial wall of the hemisphere in humans and laboratory primates. Converging evidence indicates that this region of the frontal lobe contains multiple areas involved in motor control. In the monkey, the medial wall contains four premotor areas that project directly to both the primary motor cortex and the spinal cord. These are the supplementary motor area (SMA) on the superior frontal gyrus and three motor areas buried within the cingulate sulcus. In addition, there is evidence that a fifth motor field. the pre-SMA. lies rostral to the SMA proper. Recent physiological observations provide evidence for functional differences among these motor fields.
In the human, no consensus exists on the number of distinct motor fields on the medial wall. In this review, we summarize the results of positron emission tomography (PET) studies that examined functional activation on the medial wall of humans. Our analysis suggests that it is possible to identify at least four separate cortical areas on the medial wall. Each area appears to be relatively more involved in some aspects of motor behavior than others. These cortical areas in the human appear to be analogous to the pre-SMA, the SMA proper, and two of the cingulate motor areas of the monkey. We believe that these correspondences and the anatomical framework we describe will be important for unraveling the motor functions of the medial wall of the hemisphere.
The cytoarchitecture of the human and the macaque monkey dorsolateral prefrontal cortex has been examined in a strictly comparative manner in order to resolve major discrepancies between the available segmentations of this cortical region in the human and the monkey brain. In addition, the connections of the dorsolateral prefrontal cortical areas were re-examined in the monkey. The present analysis showed that only a restricted portion of what had previously been labelled as area 46 in the monkey has the same characteristics as area 46 of the human brain; the remaining part of this monkey region has the characteristics of a portion of the middle frontal gyrus in the human brain that had previously been included as part of area 9. We have labelled this cortical area as 9/46 in both species. These two areas (i.e. 46 and 9/46), which constitute the lower half of the mid-dorsolateral frontal cortex, have a well-developed granular layer IV, and can easily be distinguished from area 9, on the upper part of the mid-dorsolateral region, which does not have a well-developed granular layer IV. Area 9 has the same basic pattern of connections as areas 46 and 9/46, but, unlike the latter areas, it does not receive input from the lateral parietal cortex. Caudal to area 9, on the dorsomedial portion of the frontal cortex, there is a distinct strip of cortex (area 8B) which, unlike area 9, receives significant input from the prestriate cortex and the medial parietal cortex. The present results provide a basis for a closer integration of findings from functional neuroimaging studies in human subjects with experimental work in the monkey.
We found that medial frontal cortex activity associated with action monitoring (detecting errors and behavioral conflict) depended on activity in the lateral prefrontal cortex. We recorded the error-related negativity (ERN), an event-related brain potential proposed to reflect anterior cingulate action monitoring, from individuals with lateral prefrontal damage or age-matched or young control participants. In controls, error trials generated greater ERN activity than correct trials. In individuals with lateral prefrontal damage, however, correct-trial ERN activity was equal to error-trial ERN activity. Lateral prefrontal damage also affected corrective behavior. Thus the lateral prefrontal cortex seemed to interact with the anterior cingulate cortex in monitoring behavior and in guiding compensatory systems.
A database of positron-emission-tomography studies published between January 1993 and November 1996 was created to address several questions regarding the function and connectivity of the human anterior cingulate cortex (ACC). Using this database, we have previously reported on the relationship between behavioural variables and the probability of blood-flow response in distinct subdivisions of the ACC. The goal of the current analysis was to discover which areas of the frontal cortex show increased blood-flow co-occurring consistently with increased blood-flow in the ACC. Analyses of the frequency distributions of peaks in the ACC and the remaining frontal cortex (FC) yielded several important findings. First, FC peaks in the precentral gyrus, superior frontal gyrus, middle frontal gyrus, inferior frontal gyrus, medial frontal gyrus and orbitomedial frontal gyri were more frequent in subtractions that also yielded a peak in the ACC than in those that did not yield an ACC peak. Second, regional differences in the frequency distribution of these FC peaks were observed when the ACC peaks were subdivided into the rostral versus caudal ACC and supracallosal versus subcallosal ACC. Peaks in the precentral gyrus and in the vicinity of the supplementary motor area were more prevalent in subtractions with co-occurring peaks in the caudal than with the rostral ACC. Peaks in the middle frontal gyrus were more frequent in subtractions with co-occurring peaks in the paralimbic part of the supracallosal ACC, relative to the subcallosal or limbic supracallosal ACC. These observations are consistent with known differences in the anatomic connectivity in these cortical regions, as defined in non-human primates. Further analyses of the influence of behavioural variables on the relationships between the ACC and other regions of the frontal cortex suggested that this type of meta-analysis may provide testable hypotheses about functional and effective connectivity within the human frontal lobe.
Controversy surrounds the function of the anterior cingulate cortex. Recent discussions about its role in behavioural control have centred on three main issues: its involvement in motor control, its proposed role in cognition and its relationship with the arousal/drive state of the organism. I argue that the overlap of these three domains is key to distinguishing the anterior cingulate cortex from other frontal regions, placing it in a unique position to translate intentions to actions.
A neglected question regarding cognitive control is how control processes might detect situations calling for their involvement. The authors propose here that the demand for control may be evaluated in part by monitoring for conflicts in information processing. This hypothesis is supported by data concerning the anterior cingulate cortex, a brain area involved in cognitive control, which also appears to respond to the occurrence of conflict. The present article reports two computational modeling studies, serving to articulate the conflict monitoring hypothesis and examine its implications. The first study tests the sufficiency of the hypothesis to account for brain activation data, applying a measure of conflict to existing models of tasks shown to engage the anterior cingulate. The second study implements a feedback loop connecting conflict monitoring to cognitive control, using this to simulate a number of important behavioral phenomena.
Reward processing involves both appetitive and consummatory phases. We sought to examine whether reward anticipation vs outcomes would recruit different regions of ventral forebrain circuitry using event-related fMRI. Nine healthy volunteers participated in a monetary incentive delays task in which they either responded to a cued target for monetary reward, responded to a cued target for no reward, or did not respond to a cued target during scanning. Multiple regression analyses indicated that while anticipation of reward vs non-reward activated foci in the ventral striatum, reward vs non-reward outcomes activated foci in the ventromedial frontal cortex. These findings suggest that reward anticipation and outcomes may differentially recruit distinct regions that lie along the trajectory of ascending dopamine projections.
As monkeys perform schedules containing several trials with a visual cue indicating reward proximity, their error rates decrease as the number of remaining trials decreases, suggesting that their motivation and/or reward expectancy increases as the reward approaches. About one-third of single neurons recorded in the anterior cingulate cortex of monkeys during these reward schedules had responses that progressively changed strength with reward expectancy, an effect that disappeared when the cue was random. Alterations of this progression could be the basis for the changes from normal that are reported in anterior cingulate population activity for obsessive-compulsive disorder and drug abuse, conditions characterized by disturbances in reward expectancy.
Recent neurophysiological studies reveal that neurons in certain brain structures carry specific signals about past and future rewards. Dopamine neurons display a short-latency, phasic reward signal indicating the difference between actual and predicted rewards. The signal is useful for enhancing neuronal processing and learning behavioral reactions. It is distinctly different from dopamine's tonic enabling of numerous behavioral processes. Neurons in the striatum, frontal cortex, and amygdala also process reward information but provide more differentiated information for identifying and anticipating rewards and organizing goal-directed behavior. The different reward signals have complementary functions, and the optimal use of rewards in voluntary behavior would benefit from interactions between the signals. Addictive psychostimulant drugs may exert their action by amplifying the dopamine reward signal.
Previous studies have reported electrophysiological brain activity that is modulated when subjects commit errors in speeded reaction time tasks. This activity is thought to index an action monitoring system in anterior cingulate cortex that signals the need for performance adjustments to minimize the risk of future errors. Consistent with this view, we report here that performance errors are foreshadowed in a modulation of this brain activity on the immediately preceding trial. We propose that this modulation reflects fluctuations in the efficiency of the action monitoring system, which may occasionally compromise subsequent performance and thus comprise a prelude to performance errors.
Prefrontal cortex (PFC) supports flexible behavior by mediating cognitive control, though the elemental forms of control supported by PFC remain a central debate. Dorsolateral PFC (DLPFC) is thought to guide response selection under conditions of response conflict or, alternatively, may refresh recently active representations within working memory. Lateral frontopolar cortex (FPC) may also adjudicate response conflict, though others propose that FPC supports higher order control processes such as subgoaling and integration. Anterior cingulate cortex (ACC) is hypothesized to upregulate response selection by detecting response conflict; it remains unclear whether ACC functions generalize beyond monitoring response conflict. The present fMRI experiment directly tested these competing theories regarding the functional roles of DLPFC, FPC, and ACC. Results reveal dissociable control processes in PFC, with mid-DLPFC selectively mediating resolution of response conflict and FPC further mediating subgoaling/integration. ACC demonstrated a broad sensitivity to control demands, suggesting a generalized role in modulating cognitive control.
When there is conflict regarding which action plan to execute in response to a sensory stimulus, the brain exerts greater cognitive control to try and resolve the conflict. In their Perspective, [Matsumoto and Tanaka][1] explain new work ([ Kerns et al .][2]) showing that the detection of action-plan conflict by the anterior cingulate cortex leads to recruitment of greater cognitive control in the lateral prefrontal cortex.
[1]: http://www.sciencemag.org/cgi/content/full/303/5660/969
[2]: http://www.sciencemag.org/cgi/content/short/303/5660/1023
The error-related negativity (ERN) is an event-related brain potential elicited by error commission and by presentation of feedback stimuli indicating incorrect performance. In this study, the authors report two experiments in which participants tried to learn to select between response options by trial and error, using feedback stimuli indicating monetary gains and losses. The results demonstrate that the amplitude of the ERN is determined by the value of the eliciting outcome relative to the range of outcomes possible, rather than by the objective value of the outcome. This result is discussed in terms of a recent theory that holds that the ERN reflects a reward prediction error signal associated with a neural system for reinforcement learning.