Franziska M. Korb

Post-Error Behavioral Adjustments Are Facilitated by Activation and Suppression of Task-Relevant and Task-Irrelevant Information Processing

Error monitoring by the posterior medial frontal cortex (pMFC) has been linked to post-error behavioral adaptation effects and cognitive control dynamics in lateral prefrontal cortex (LPFC). It remains unknown, however, whether control adjustments following errors produce post-error behavioral adjustments (PEBAs) by inhibiting inappropriate responses or facilitating goal-directed ones. Here we used functional magnetic resonance imaging to investigate the hemodynamic correlates of PEBAs in a stimulus–response compatibility task. Our task was designed to test whether PEBAs are implemented by suppressing motor responses primed by irrelevant stimulus features (face location), redirecting attention to relevant features (face gender), or both or neither of these possibilities. Independent of PEBAs, error-related pMFC activation was followed by post-error recruitment of prefrontal and parietal control regions and, crucially, both (1) suppressed response-related activity in sensorimotor cortex and (2) enhanced target processing in face-sensitive sensory cortex (“fusiform face area”). More importantly, by investigating the covariation between post-error hemodynamic activity and individual differences in PEBAs, we showed that modulation of task-related motor and sensory processing was dependent on whether participants produced generally slower responses (“post-error slowing”; PES) or selectively reduced interference effects (“post-error reduction of interference”; PERI), respectively. Each of these behaviorally dependent effects was mediated by distinct LPFC control mechanisms (PES: inferior frontal junction; PERI: superior frontal sulcus). While establishing relationships between PEBAs and cognitive control, our findings suggest that the neural architecture underlying sequential behavioral adaptation may be determined primarily by how control is executed by the individual when adjustments are needed.

Priming of Control: Implicit Contextual Cuing of Top-down Attentional Set

Cognitive models have long distinguished between “automatic” associative processes that can be triggered in a bottom-up fashion, and “controlled” processes, where internal goals guide information processing in a deliberate, top-down manner. However, recent behavioral studies have cast doubt on the validity of this dichotomy, showing that implicit contextual cues can modulate performance in a way suggestive of an associative triggering of specific top-down control states. Here, we harnessed functional magnetic resonance imaging in humans to test whether these behavioral findings truly reflect online, bottom-up priming of top-down attentional control settings. Using a flanker interference task where stimulus location cued the likelihood of incongruent trials, we found that the behavioral phenomenon of implicit, context-specific improvements in interference resolution was mirrored in hemodynamic activity in the medial superior parietal lobule (mSPL), previously implicated in voluntary (as opposed to primed) attention shifts. Moreover, the mSPL displayed context-specific functional coupling with visual regions involved in processing the flanker stimuli, and the modulation of the latter was predictive of the behavioral effects. Finally, the implementation of this contextual control was “on the fly,” that is, it was primed online by a switch to the context associated with high conflict. These results suggest that top-down control states can be bound into episodic event representations and can subsequently be primed by other features of those representations. Together, our findings illustrate a more intimate link between associative and controlled processing than is traditionally assumed, and place the neural substrate of that linkage in the posterior parietal cortex.

Affective Modulation of Cognitive Control is Determined by Performance-Contingency and Mediated by Ventromedial Prefrontal and Cingulate Cortex

Cognitive control requires a fine balance between stability, the protection of an on-going task-set, and flexibility, the ability to update a task-set in line with changing contingencies. It is thought that emotional processing modulates this balance, but results have been equivocal regarding the direction of this modulation. Here, we tested the hypothesis that a crucial determinant of this modulation is whether affective stimuli represent performance-contingent or task-irrelevant signals. Combining functional magnetic resonance imaging with a conflict task-switching paradigm, we contrasted the effects of presenting negative- and positive-valence pictures on the stability/flexibility trade-off in humans, depending on whether picture presentation was contingent on behavioral performance. Both the behavioral and neural expressions of cognitive control were modulated by stimulus valence and performance contingency: in the performance-contingent condition, cognitive flexibility was enhanced following positive pictures, whereas in the nonperformance-contingent condition, positive stimuli promoted cognitive stability. The imaging data showed that, as anticipated, the stability/flexibility trade-off per se was reflected in differential recruitment of dorsolateral frontoparietal and striatal regions. In contrast, the affective modulation of stability/flexibility shifts was mirrored, unexpectedly, by neural responses in ventromedial prefrontal and posterior cingulate cortices, core nodes of the “default mode” network. Our results demonstrate that the affective modulation of cognitive control depends on the performance contingency of the affect-inducing stimuli, and they document medial default mode regions to mediate the flexibility-promoting effects of performance-contingent positive affect, thus extending recent work that recasts these regions as serving a key role in on-task control processes.

Transcranial Magnetic Stimulation Intensities in Cognitive Paradigms

Background Transcranial magnetic stimulation (TMS) has become an important experimental tool for exploring the brains functional anatomy. As TMS interferes with neural activity, the hypothetical function of the stimulated area can thus be tested. One unresolved methodological issue in TMS experiments is the question of how to adequately calibrate stimulation intensities. The motor threshold (MT) is often taken as a reference for individually adapted stimulation intensities in TMS experiments, even if they do not involve the motor system. The aim of the present study was to evaluate whether it is reasonable to adjust stimulation intensities in each subject to the individual MT if prefrontal regions are stimulated prior to the performance of a cognitive paradigm. Methods and Findings Repetitive TMS (rTMS) was applied prior to a working memory task, either at the ‘fixed’ intensity of 40% maximum stimulator output (MSO), or individually adapted at 90% of the subjects MT. Stimulation was applied to a target region in the left posterior middle frontal gyrus (pMFG), as indicated by a functional magnetic resonance imaging (fMRI) localizer acquired beforehand, or to a control site (vertex). Results show that MT predicted the effect size after stimulating subjects with the fixed intensity (i.e., subjects with a low MT showed a greater behavioral effect). Nevertheless, the individual adaptation of intensities did not lead to stable effects. Conclusion Therefore, we suggest assessing MT and account for it as a measure for general cortical TMS susceptibility, even if TMS is applied outside the motor domain.

The role of anterior cingulate cortex in the affective evaluation of conflict

An influential theory of ACC function argues that this brain region plays a crucial role in the affective evaluation of performance monitoring and control demands. Specifically, control-demanding processes such as response conflict are thought to be registered as aversive signals by ACC, which in turn triggers processing adjustments to support avoidance learning. In support of conflict being treated as an aversive event, recent behavioral studies demonstrated that incongruent (i.e., conflict inducing), relative to congruent, stimuli can speed up subsequent negative, relative to positive, affective picture processing. Here, we used fMRI to investigate directly whether ACC activity in response to negative versus positive pictures is modulated by preceding control demands, consisting of conflict and task-switching conditions. The results show that negative, relative to positive, pictures elicited higher ACC activation after congruent, relative to incongruent, trials, suggesting that ACCs response to negative (positive) pictures was indeed affectively primed by incongruent (congruent) trials. Interestingly, this pattern of results was observed on task repetitions but disappeared on task alternations. This study supports the proposal that conflict induces negative affect and is the first to show that this affective signal is reflected in ACC activation.

Levels of integration in cognitive control and sequence processing in the prefrontal cortex

Cognitive control is necessary to flexibly act in changing environments. Sequence processing is needed in language comprehension to build the syntactic structure in sentences. Functional imaging studies suggest that sequence processing engages the left ventrolateral prefrontal cortex (PFC). In contrast, cognitive control processes additionally recruit bilateral rostral lateral PFC regions. The present study aimed to investigate these two types of processes in one experimental paradigm. Sequence processing was manipulated using two different sequencing rules varying in complexity. Cognitive control was varied with different cue-sets that determined the choice of a sequencing rule. Univariate analyses revealed distinct PFC regions for the two types of processing (i.e. sequence processing: left ventrolateral PFC and cognitive control processing: bilateral dorsolateral and rostral PFC). Moreover, in a common brain network (including left lateral PFC and intraparietal sulcus) no interaction between sequence and cognitive control processing was observed. In contrast, a multivariate pattern analysis revealed an interaction of sequence and cognitive control processing, such that voxels in left lateral PFC and parietal cortex showed different tuning functions for tasks involving different sequencing and cognitive control demands. These results suggest that the difference between the process of rule selection (i.e. cognitive control) and the process of rule-based sequencing (i.e. sequence processing) find their neuronal underpinnings in distinct activation patterns in lateral PFC. Moreover, the combination of rule selection and rule sequencing can shape the response of neurons in lateral PFC and parietal cortex.

Model-based analysis of context-specific cognitive control.

Interference resolution is improved for stimuli presented in contexts (e.g., locations) associated with frequent conflict. This phenomenon, the context-specific proportion congruent (CSPC) effect, has challenged the traditional juxtaposition of “automatic” and “controlled” processing because it suggests that contextual cues can prime top-down control settings in a bottom-up manner. We recently obtained support for this “priming of control” hypothesis with functional magnetic resonance imaging by showing that CSPC effects are mediated by contextually cued adjustments in processing selectivity. However, an equally plausible explanation is that CSPC effects reflect adjustments in response caution triggered by expectancy violations (i.e., prediction errors) when encountering rare events as compared to common ones (e.g., incongruent trials in a task context associated with infrequent conflict). Here, we applied a quantitative model of choice, the linear ballistic accumulator (LBA), to distil the reaction time and accuracy data from four independent samples that performed a modified flanker task into latent variables representing the psychological processes underlying task-related decision making. We contrasted models which differentially accounted for CSPC effects as arising either from contextually cued shifts in the rate of sensory evidence accumulation (“drift” models) or in the amount of evidence required to reach a decision (“threshold” models). For the majority of the participants, the LBA ascribed CSPC effects to increases in response threshold for contextually infrequent trial types (e.g., congruent trials in the frequent conflict context), suggesting that the phenomenon may reflect more a prediction error-triggered shift in decision criterion rather than enhanced sensory evidence accumulation under conditions of frequent conflict.

Dissociable causal roles for left and right parietal cortex in controlling attentional biases from the contents of working memory

The contents of working memory (WM) steer visual attention, but the extent of this guidance can be strategically enhanced or inhibited when WM content is reliably helpful or harmful to a visual task. Current understanding of the neural substrates mediating the cognitive control over WM biases is limited, however, by the correlational nature of functional MRI approaches. A recent fMRI study provided suggestive evidence for a functional lateralization of these control processes in posterior parietal cortex (PPC): activity in left PPC correlated with the presentation of WM cues that ought to be strategically enhanced to optimize performance, while activity in the right PPC correlated with the presentation of cues that ought to be inhibited to prevent detrimental attentional biases in a visual search. Here, we aimed to directly assess whether the left and right PPC are causally involved in the cognitive control of WM biases, and to clarify their precise functional contributions. We therefore applied 1 Hz repetitive transcranial magnetic stimulation (rTMS) to left and right PPC (and a vertex control site) prior to administering a behavioral task assessing WM biasing control functions. We observed that the perturbation of left PPC eliminated the strategic benefit of predictably helpful WM cueing, while the perturbation of right PPC amplified the cost of unpredictable detrimental WM cueing. The left and right PPC thus play distinct causal roles in WM-attention interactions: the left PPC to maximize benefits, and the right PPC to minimize costs, of internally maintained content on visual attention.

Comparing neural substrates of emotional vs. non-emotional conflict modulation by global control context.

The efficiency with which the brain resolves conflict in information processing is determined by contextual factors that modulate internal control states, such as the recent (local) and longer-term (global) occurrence of conflict. Local “control context” effects can be observed in trial-by-trial adjustments to conflict (congruency sequence effects: less interference following incongruent trials), whereas global control context effects are reflected in adjustments to the frequency of conflict encountered over longer sequences of trials (“proportion congruent effects”: less interference when incongruent trials are frequent). Previous neuroimaging and lesion studies suggest that the modulation of conflict-control processes by local control context relies on partly dissociable neural circuits for cognitive (non-emotional) vs. emotional conflicts. By contrast, emotional and non-emotional conflict-control processes have not been contrasted with respect to their modulation by global control context. We addressed this aim in a functional magnetic resonance imaging (fMRI) study that varied the proportion of congruent trials in emotional vs. non-emotional conflict tasks across blocks. We observed domain-general conflict-related signals in the dorsal anterior cingulate cortex (dACC) and pre-supplementary motor area and, more importantly, task-domain also interacted with global control context effects: specifically, the dorsal striatum and anterior insula tracked control-modulated conflict effects exclusively in the emotional domain. These results suggest that, similar to the neural mechanisms of local control context effects, there are both overlapping as well as distinct neural substrates involved in the modulation of emotional and non-emotional conflict-control by global control context.

Reduced Risk-Taking following Disruption of the Intraparietal Sulcus

Decision makers frequently encounter opportunities to pursue great gains—assuming they are willing to accept greater risks. Previous neuroimaging studies have shown that activity in the intraparietal sulcus (IPS) and the inferior frontal junction (IFJ) are associated with individual preferences for economic risk (“known unknowns,” e.g., a 50% chance of winning