The neural basis of error detection: conflict monitoring and the error-related negativity

According to a recent theory, anterior cingulate cortex is sensitive to response conflict, the coactivation of mutually incompatible responses. The present research develops this theory to provide a new account of the error-related negativity (ERN), a scalp potential observed following errors. Connectionist simulations of response conflict in an attentional task demonstrated that the ERN--its timing and sensitivity to task parameters--can be explained in terms of the conflict theory. A new experiment confirmed predictions of this theory regarding the ERN and a second scalp potential, the N2, that is proposed to reflect conflict monitoring on correct response trials. Further analysis of the simulation data indicated that errors can be detected reliably on the basis of post-error conflict. It is concluded that the ERN can be explained in terms of response conflict and that monitoring for conflict may provide a simple mechanism for detecting errors.     

A computational account of altered error processing in older age: Dopamine and the error-related negativity

When participants commit errors or receive feedback signaling that they have made an error, a negative brain potential is elicited. According to Holroyd and Coles’s (in press) neurocomputational model of error processing, this error-related negativity (ERN) is elicited when the brain first detects that the consequences of an action are worse than expected. To study age-related changes in error processing, we obtained performance and ERN measures of younger and high-functioning older adults. Experiment 1 demonstrated reduced ERN amplitudes in older adults in the context of otherwise intact brain potentials. This result could not be attributed to uncertainty about the required response in older adults. Experiment 2 revealed impaired performance and reduced response- and feedback-related ERNs of older adults in a probabilistic learning task. These age changes could be simulated by manipulation of a single parameter of the neurocomputational model, this manipulation corresponding to weakened phasic activity of the mesencephalic dopamine system.     

错误加工意识水平与错误相关脑电成分

错误加工的意识水平包括有意识水平、无意识水平以及介于两者之间的意识水平。意识水平的划分通常通过错误报告范式实现。根据其对意识水平的划分, 可以将相关研究分为二元意识水平研究和多元意识水平研究。已有研究, 主要关注于意识水平与错误诱发的Ne、Pe的关系。两类研究一致表明, Pe受意识水平的调节, 但是并没有得到fMRI研究的一致支持。对Ne的激活是否独立于意识, 研究者们仍存在争议。还有研究考察了注意对错误意识水平的影响, 结果支持Pe与意识水平共变的观点, 但是对注意如何影响错误意识水平还存在质疑。在对以上研究进行整理与分析的基础上, 对将来的研究提出了建议。     

Externalizing psychopathology and the error-related negativity

Prior research has demonstrated that antisocial behavior, substance-use disorders, and personality dimensions of aggression and impulsivity are indicators of a highly heritable underlying dimension of risk, labeled externalizing. Other work has shown that individual trait constructs within this psychopathology spectrum are associated with reduced self-monitoring, as reflected by amplitude of the error-related negativity (ERN) brain response. In this study of undergraduate subjects, reduced ERN amplitude was associated with higher scores on a self-report measure of the broad externalizing construct that links these various indicators. In addition, the ERN was associated with a response-locked increase in anterior theta (4-7 Hz) oscillation; like the ERN, this theta response to errors was reduced among high-externalizing individuals. These findings suggest that neurobiologically based deficits in endogenous action monitoring may underlie generalized risk for an array of impulse-control problems.     

Errors are aversive: defensive motivation and the error-related negativity

The error-related negativity (ERN) is a negative deflection in the event-related potential that is maximal approximately 50 ms after the commission of an error. The ERN is generated in the anterior cingulate cortex, a region of the medial prefrontal cortex implicated in both cognitive and emotional processing. Affective and motivational variables influence the magnitude of the ERN, which suggests that the ERN may relate to emotional or motivational aspects of error detection. In the present study, we evaluated the possibility that errors prime defensive motivational responses, and that the ERN may predict the magnitude of defensive reactivity after errors. We found that (a) the defensive startle response was larger following errors than following correct responses, and (b) the magnitude of the ERN predicted the degree of startle potentiation following errors. Thus, response errors prime defensive motivation-and the ERN predicts individual differences in the aversive response to errors.     

Response monitoring, the error-related negativity, and differences in social behavior in autism

Children with autism not only display social impairments but also significant individual differences in social development. Understanding the source of these differences, as well as the nature of social impairments, is important for improved diagnosis and treatments for these children. Current theory and research suggests that individual differences in response monitoring, a specific function of the anterior cingulate cortex (ACC), may contribute to social-emotional and social-cognitive impairments and individual differences in autism. To examine this hypothesis, we used a modified flanker task to assess an ERP index of response monitoring, the error-related negativity (ERN), in a sample of higher function children with autism (HFA) and an IQ-matched control sample. The results revealed a significant Diagnostic group by Verbal IQ interaction on ERN amplitude indicating that the most verbally capable HFA children displayed significantly larger ERN amplitudes than did the control children. Within the HFA sample, greater ERN amplitude was also related to parent reports of fewer symptoms of social interaction impairments, fewer internalizing problems, but more externalizing problems, although these associations were reduced to nonsignificance when medication status was controlled. The latter results complement previous observations from imaging studies of a significant association between ACC activity and social symptoms and impairments in autism. The implications of these results for future research on brain-behavior relations, as well as treatment related research with children with autism are discussed.     

The effect of trial-to-trial feedback on the error-related negativity and its relationship with anxiety

Individuals with anxiety disorders and related personality traits are characterized by increased error-related brain activity, as measured by the error-related negativity (ERN) in simple speeded response tasks. An absent, or opposite, relation between anxiety and the ERN has been reported in studies that employed reinforcement learning paradigms with trial-to-trial feedback. Understanding the effect of trial-to-trial feedback on the ERN may help clarify these results and can further elucidate the impact of feedback on performance monitoring. In the present study, 30 undergraduate participants performed two versions of the arrowhead version of the flanker task in counterbalanced order: one with trial-to-trial feedback and one without. The participants were slower and more accurate in the task with trial-to-trial feedback; however, the ERN was equivalent between the two tasks. Larger ERNs were related to higher trait anxiety, but only in the version without trial-to-trial feedback. These findings show that although trial-to-trial performance feedback impacts behavioral measures, it does not affect the ERN; moreover, the presence of trial-to-trial feedback moderates the relationship between the ERN and anxiety.     

The neural basis of haptic object processing.

We review the organization of the neural networks that underlie haptic object processing and compare that organization with the visual system. Haptic object processing is separated into at least two neural pathways, one for geometric properties or shape, and one for material properties, including texture. Like vision, haptic processing pathways are organized into a hierarchy of processing stages, with different stages represented by different brain areas. In addition, the haptic pathway for shape processing may be further subdivided into different streams for action and perception. These streams may be analogous to the action and perception streams of the visual system and represent two points of neural convergence for vision and haptics.     

Individual differences in working memory capacity predict action monitoring and the error-related negativity

Neuroscience suggests that the anterior cingulate cortex (ACC) is responsible for conflict monitoring and the detection of errors in cognitive tasks, thereby contributing to the implementation of attentional control. Though individual differences in frontally mediated goal maintenance have clearly been shown to influence outward behavior in interference-rich contexts, it is unclear whether corresponding differences exist in neural responses that arise out of the ACC. To investigate this possibility, we conducted an electrophysiological study using a variant of the Simon Task, recording event-related potentials (ERPs) in healthy normal individuals with varying working memory capacity (high vs. low spans; a behavioral proxy for variability in goal maintenance). Primary analyses focused on the magnitude of the error-related negativity (ERN), a response-locked ERP component associated with the commission of errors thought to arise because of action monitoring in the ACC. Our results revealed that frontally mediated working memory capacity may alter error monitoring by the ACC, with high spans showing a greater ERN than low spans. These individual differences were also observed in the posterror positivity, a response-locked ERP component associated with updating cognitive strategies, suggesting greater awareness of errors with increased working memory capacity. These results are interpreted within 2-process models of attentional control, suggesting individuals with greater working memory capacity may better maintain task goals by more strongly biasing neural activity in frontal-executive networks.     

Student learning as the basis for reinforcement to the instructor

The present study reinforced the two instructors of two students with profound mental retardation first for the instructors' training behaviors, then for improved student learning in a multiple baseline across responses design. Reinforcement of the instructors' training behaviors increased their training behaviors, replicating the findings of previous studies, but had no effect on student learning. When the reinforcement for the instructors was instead made contingent upon student learning, the student learning improved and the instructors' high level of training behaviors was maintained. In addition, the instructors began to request training for themselves. These findings suggest that staff motivation programs for instructors might usefully base the reinforcers not only on the instructor's performance, but on the progress of their students.     

语言认知老化机制及其神经基础

老龄化个体出现的认知障碍在某种程度上是由于语言的认知老化所导致,因此,了解语言认知老化的具体表现、老化机制及其神经基础对于延缓语言认知老化、矫正老年痴呆有重要意义。首先基于理解和产生两个维度,分别从词汇、句子和文本三个层面分析了语言认知老化的表现,而后着重从工作记忆的角度分析了语言认知老化的认知机制,在此基础上,进一步从词汇加工、语义提取、句法分析三个方面分析了语言认知老化的神经基础,并指出半球不对称性减弱和脑区的弥散性激活是语言认知老化的主要神经表现。最后,围绕着语言认知老化的"发展进程、领域表现形式、干预矫正方法"三个必须解决的问题展开讨论,为该领域未来的发展做出理论上的铺垫。     

The impact of cognitive deficits on conflict monitoring. Predictable dissociations between the error-related negativity and N2

Monitoring of ongoing processing plays a critical role in regulating cognitive function. Two event-related potential components, the error-related negativity (ERN) and N2, have been proposed to reflect this monitoring function. Specifically, it has been suggested that both components reflect the role of anterior cingulate cortex (ACC) in monitoring for the occurrence of response conflict. This view appears to be challenged by findings that alcohol consumption and lesions in ACC have dissociable effects on the ERN and N2. Using a computational model, the present research demonstrates that the conflict-monitoring theory can account for these dissociations in terms of the dissociable effects of alcohol and ACC lesions on processing of relevant stimulus information (which determines ERN amplitude) and processing of irrelevant, distracting information (which determines N2 amplitude). The simulation results suggest new interpretations of the cognitive deficits caused by alcohol consumption (impaired stimulus processing) and ACC lesions (impaired attentional control).     

Hierarchically organized behavior and its neural foundations: a reinforcement learning perspective

Research on human and animal behavior has long emphasized its hierarchical structure—the divisibility of ongoing behavior into discrete tasks, which are comprised of subtask sequences, which in turn are built of simple actions. The hierarchical structure of behavior has also been of enduring interest within neuroscience, where it has been widely considered to reflect prefrontal cortical functions. In this paper, we reexamine behavioral hierarchy and its neural substrates from the point of view of recent developments in computational reinforcement learning. Specifically, we consider a set of approaches known collectively as hierarchical reinforcement learning, which extend the reinforcement learning paradigm by allowing the learning agent to aggregate actions into reusable subroutines or skills. A close look at the components of hierarchical reinforcement learning suggests how they might map onto neural structures, in particular regions within the dorsolateral and orbital prefrontal cortex. It also suggests specific ways in which hierarchical reinforcement learning might provide a complement to existing psychological models of hierarchically structured behavior. A particularly important question that hierarchical reinforcement learning brings to the fore is that of how learning identifies new action routines that are likely to provide useful building blocks in solving a wide range of future problems. Here and at many other points, hierarchical reinforcement learning offers an appealing framework for investigating the computational and neural underpinnings of hierarchically structured behavior.     

Towards a neural basis of auditory sentence processing

Functional dissociations within the neural basis of auditory sentence processing are difficult to specify because phonological, syntactic and semantic information are all involved when sentences are perceived. In this review I argue that sentence processing is supported by a temporo-frontal network. Within this network, temporal regions subserve aspects of identification and frontal regions the building of syntactic and semantic relations. Temporal analyses of brain activation within this network support syntax-first models because they reveal that building of syntactic structure precedes semantic processes and that these interact only during a later stage.     

Backward inhibitory learning in honeybees: a behavioral analysis of reinforcement processing

One class of theoretical accounts of associative learning suggests that reinforcers are processed according to learning rules that minimize the predictive error between the expected strength of future reinforcement and its actual strength. The omission of reinforcement in a situation where it is expected leads to inhibitory learning of stimuli indicative for such a violation of the prediction. There are, however, results indicating that inhibitory learning can also be induced by other mechanisms. Here, we present data from olfactory reward conditioning in honeybees that show that (1) one- and multiple-trial backward conditioning results in conditioned inhibition (CI); (2) the inhibition is maximal for a 15-sec interval between US and CS; (3) there is a nonmonotonic dependency on the degree of CI from the US-CS interval during backward pairing; and (4) the prior association of context stimuli with reinforcement is not necessary for the development of CI. These results cannot be explained by models that only minimize a prediction error. Rather, they are consistent with models of associative learning that, in addition, assume that learning depends on the temporal overlap of a CS with two processes, a fast excitatory and a slow inhibitory one, both evoked by a reinforcer. The fmdings from this behavioral analysis of reinforcement processing are compared with the known properties of an individual, identified neuron involved in reinforcement processing in the bee brain, to further understand the mechanisms underlying predictive reward learning.     

Adolescent-specific patterns of behavior and neural activity during social reinforcement learning

Humans are sophisticated social beings. Social cues from others are exceptionally salient, particularly during adolescence. Understanding how adolescents interpret and learn from variable social signals can provide insight into the observed shift in social sensitivity during this period. The present study tested 120 participants between the ages of 8 and 25 years on a social reinforcement learning task where the probability of receiving positive social feedback was parametrically manipulated. Seventy-eight of these participants completed the task during fMRI scanning. Modeling trial-by-trial learning, children and adults showed higher positive learning rates than did adolescents, suggesting that adolescents demonstrated less differentiation in their reaction times for peers who provided more positive feedback. Forming expectations about receiving positive social reinforcement correlated with neural activity within the medial prefrontal cortex and ventral striatum across age. Adolescents, unlike children and adults, showed greater insular activity during positive prediction error learning and increased activity in the supplementary motor cortex and the putamen when receiving positive social feedback regardless of the expected outcome, suggesting that peer approval may motivate adolescents toward action. While different amounts of positive social reinforcement enhanced learning in children and adults, all positive social reinforcement equally motivated adolescents. Together, these findings indicate that sensitivity to peer approval during adolescence goes beyond simple reinforcement theory accounts and suggest possible explanations for how peers may motivate adolescent behavior.