Nicola K. Ferdinand

The Processing of Unexpected Positive Response Outcomes in the Mediofrontal Cortex

The human mediofrontal cortex, especially the anterior cingulate cortex, is commonly assumed to contribute to higher cognitive functions like performance monitoring. How exactly this is achieved is currently the subject of lively debate but there is evidence that an events valence and its expectancy play important roles. One prominent theory, the reinforcement learning theory by Holroyd and colleagues (2002, 2008), assigns a special role to feedback valence, while the prediction of response–outcome (PRO) model by Alexander and Brown (2010, 2011) claims that the mediofrontal cortex is sensitive to unexpected events regardless of their valence. However, paradigms examining this issue have included confounds that fail to separate valence and expectancy. In the present study, we tested the two competing theories of performance monitoring by using an experimental task that separates valence and unexpectedness of performance feedback. The feedback-related negativity of the event-related potential, which is commonly assumed to be a reflection of mediofrontal cortex activity, was elicited not only by unexpected negative feedback, but also by unexpected positive feedback. This implies that the mediofrontal cortex is sensitive to the unexpectedness of events in general rather than their valence and by this supports the PRO model.

Error and deviance processing in implicit and explicit sequence learning

In this experiment, we examined the extent to which error-driven learning may operate under implicit learning conditions. We compared error monitoring in a sequence learning task in which stimuli consisted of regular, irregular, or random sequences. Subjects were either informed (explicit condition) or not informed (implicit condition) about the existence of the sequence. For both conditions, reaction times were faster to stimuli from regular sequences than from random sequences, thus supporting the view that sequence learning occurs irrespective of learning condition. Response-locked event-related potentials (ERPs) showed a pronounced ERN/Ne, thereby signaling the detection of committed errors. Deviant stimuli from irregular sequences elicited an N2b component that developed in the course of the experiment, albeit faster for explicit than implicit learners. This observation supports the view that deviant events acquire the status of perceived errors during explicit and implicit learning, and thus, an N2b is generated resembling the ERN/Ne to committed errors. While performing the task, expectations about upcoming events are generated, compared to the actual events, and evaluated on the dimension better or worse than expected. The accuracy of this process improves with learning, as shown by a gradual increase in N2b amplitude as a function of learning. Additionally, a P3b, which is thought to mirror conscious processing of deviant stimuli and is related to updating of working memory representations, was found for explicit learners only.

Timing Matters: The Impact of Immediate and Delayed Feedback on Artificial Language Learning

In the present experiment, we used event-related potentials (ERP) to investigate the role of immediate and delayed feedback in an artificial grammar learning (AGL) task. Two groups of participants were engaged in classifying non-word strings according to an underlying rule system, not known to the participants. Visual feedback was provided after each classification either immediately or with a short delay of 1 s. Both groups were able to learn the artificial grammar system as indicated by an increase in classification performance. However, the gain in performance was significantly larger for the group receiving immediate feedback as compared to the group receiving delayed feedback. Learning was accompanied by an increase in P300 activity in the ERP for delayed as compared to immediate feedback. Irrespective of feedback delay, both groups exhibited learning related decreases in the feedback-related positivity (FRP) elicited by positive feedback only. The feedback-related negativity (FRN), however, remained constant over the course of learning. These results suggest, first, that delayed feedback is less effective for AGL as task requirements are very demanding, and second, that the FRP elicited by positive prediction errors decreases with learning while the FRN to negative prediction errors is elicited in an all-or-none fashion by negative feedback throughout the entire experiment.

Age-related changes in processing positive and negative feedback: is there a positivity effect for older adults?

Older people sometimes show a bias toward the processing of positive information. In this study, we used an event-related potential approach to examine whether such a positivity bias is also present during feedback processing in older adults. Our results suggest that a fast initial monitoring process, as reflected in the feedback-related negativity (FRN), is sensitive to the expectancy of events irrespective of their valence for older (aged 70-77 years) as well as younger (aged 20-27 years) adults. In contrast, in a later evaluation process, associated with memory updating and indexed by the P300, both age groups preferably processed unexpected positive feedback. However, younger adults additionally differentiated between unexpected negative and expected feedback while older adults did not, probably due to a lower working memory capacity.

Developmental changes in performance monitoring: how electrophysiological data can enhance our understanding of error and feedback processing in childhood and adolescence.

Performance monitoring includes learning from errors and feedback and depends on the functioning of the mediofrontal cortex, especially the anterior cingulate cortex (ACC) and the mesencephalic dopamine system. Error and feedback monitoring develop during childhood until early adulthood and are important in a lot of learning situations. The aims of this article are twofold: First, to review the present literature on the development of performance monitoring, and second, to highlight how electrophysiological data can contribute to the understanding of error and feedback processing in childhood and adolescence.

Different aspects of performance feedback engage different brain areas: Disentangling valence and expectancy in feedback processing

Evaluating the positive and negative outcomes of our behaviour is important for action selection and learning. Such reinforcement learning has been shown to engage a specific neural circuitry including the mesencephalic dopamine system and its target areas, the striatum and medial frontal cortex, especially the anterior cingulate cortex (ACC). An intensively pursued debate regards the prevailing influence of feedback expectancy and feedback valence on the engagement of these two brain regions in reinforcement learning and their respective roles are far from being understood. To this end, we used a time estimation task with three different types of feedback that allows disentangling the effect of feedback valence and expectancy using functional magnetic resonance imaging (fMRI). Our results show greater ACC activation after unexpected positive and unexpected negative feedback than after expected feedback and by this sensitivity to unexpected events in general irrespective of their valence.

Age-differential effects on updating cue information: evidence from event-related potentials.

Recent models on cognitive aging consider the ability to maintain and update context information to be a key source of age-related impairments in various cognitive tasks (Braver & Barch in Neuroscience & Biobehavioral Reviews, 26: 809–817, 2002). Context updating has been investigated with a modified AX-continuous-performance task by comparing performance and brain activity between context-dependent trials (i.e., correct responses require updating of the preceding cue information) and context-independent trials (i.e., correct responses are independent of cue information). We used an event-related potential (ERP) approach to identify sources of age differences in context processing in the early and late processing of cue information. Our behavioral data showed longer latencies and higher error rates on context-dependent than on context-independent trials for older than for younger adults, suggesting age-related impairments in context updating. The ERP data revealed larger P3b amplitudes for context-dependent than for context-independent trials only in younger adults. In contrast, in older adults, P3b amplitudes were more evenly distributed across the scalp and did not differ between context conditions. Interestingly, older but not younger adults were sensitive to changes of cue identity, as indicated by larger P3b amplitudes on cue-change than on cue-repeat trials, irrespective of the actual context condition. We also found a larger CNV on context-dependent than on context-independent trials, reflecting active maintenance of context information and response preparation. The age-differential effects in the P3b suggest that both younger and older adults were engaged in updating task-relevant information, but relied on different information: Whereas younger participants indeed relied on context cues to update and reconfigure the task settings, older adults relied on changes in cue identity, irrespective of context information.

The influence of monetary incentives on context processing in younger and older adults: an event-related potential study

Recent evidence has indicated that neuronal activity related to reward anticipation benefits subsequent stimulus processing, but the effect of penalties remains largely unknown. Since the dual-mechanisms-of-control theory (DMC; Braver & Barch, Neuroscience and Biobehavioral Reviews, 26, 809–81, 2002) assumes that temporal differences in context updating underlie age differences in cognitive control, in this study we investigated whether motivational cues (signaling the chance to win or the risk to lose money, relative to neutral cues) preceding context information in a modified AX-CPT paradigm influence the temporal stages of context processing in younger and older adults. In the behavioral data, younger adults benefited from gain cues, evident in their enhanced context updating, whereas older adults exhibited slowed responding after motivational cues, irrespective of valence. Event-related potentials (ERPs) revealed that the enhanced processing of motivational cues in the P2 and P3b was mainly age-invariant, whereas age-differential effects were found for the ERP correlates of context processing. Younger adults showed improved context maintenance (i.e., a larger negative-going CNV), as well as increased conflict detection (larger N450) and resolution (indicated by a sustained positivity), whenever incorrect responding would lead to a monetary loss. In contrast, motivationally salient cues benefited context representations (in cue-locked P3b amplitudes), but increased working memory demands during response preparation (via a temporally prolonged P3b) in older adults. In sum, motivational valence and salience effects differentially modulated the temporal stages of context processing in younger and older adults. These results are discussed in terms of the DMC theory, recent findings of emotion regulation in old age, and the relationship between cognitive and affective processing.

Event-related potential correlates of declarative and non-declarative sequence knowledge

The goal of the present study was to demonstrate that declarative and non-declarative knowledge acquired in an incidental sequence learning task contributes differentially to memory retrieval and leads to dissociable ERP signatures in a recognition memory task. For this purpose, participants performed a sequence learning task and were classified as verbalizers, partial verbalizers, or nonverbalizers according to their ability to verbally report the systematic response sequence. Thereafter, ERPs were recorded in a recognition memory task time-locked to sequence triplets that were either part of the previously learned sequence or not. Although all three groups executed old sequence triplets faster than new triplets in the recognition memory task, qualitatively distinct ERP patterns were found for participants with and without reportable knowledge. Verbalizers and, to a lesser extent, partial verbalizers showed an ERP correlate of recollection for parts of the incidentally learned sequence. In contrast, nonverbalizers showed a different ERP effect with a reverse polarity that might reflect priming. This indicates that an ensemble of qualitatively different processes is at work when declarative and non-declarative sequence knowledge is retrieved. By this, our findings favor a multiple-systems view postulating that explicit and implicit learning are supported by different and functionally independent systems.

Feedback processing in children and adolescents: Is there a sensitivity for processing rewarding feedback?

Developmental studies indicate that children rely more on external feedback than adults. Some of these studies claim that they additionally show higher sensitivity toward positive feedback, while others find they preferably use negative feedback for learning. However, these studies typically did not disentangle feedback valence and expectancy, which might contribute to the controversial results. The present study aimed at examining the neurophysiological correlates of feedback processing in children (8-10 years) and adolescents (12-14 years) in a time estimation paradigm that allows separating the contribution of valence and expectancy. Our results show that in the feedback-related negativity (FRN), an event-related potential (ERP) reflecting the fast initial processing of feedback stimuli, children and adolescents did not differentiate between unexpected positive and negative feedback. Thus, they did not show higher sensitivity to positive feedback. The FRN did also not differentiate between expected and unexpected feedback, as found for adults. In contrast, in a later processing stage mirrored in the P300 component of the ERP, children and adolescents processed the feedbacks unexpectedness. Interestingly, adolescents with better behavioral adaptation (high-performers) also had a more frontal P300 expectancy effect. Thus, the recruitment of additional frontal brain regions might lead to better learning from feedback in adolescents.