Internal attention modulates the functional state of novel stimulus-response associations in working memory.

Information in working memory (WM) is crucial for guiding behavior. However, not all WM representations are equally relevant simultaneously. Current theoretical frameworks propose a functional dissociation between 'latent' and 'active' states, in which relevant representations are prioritized into an optimal (active) state to face current demands, while relevant information that is not immediately needed is maintained in a dormant (latent) state. In this context, task demands can induce rapid and flexible prioritization of information from latent to active state. Critically, these functional states have been primarily studied using simple visual memories, with attention selecting and prioritizing relevant representations to serve as templates to guide subsequent behavior. It remains unclear whether more complex WM representations, such as novel stimulus-response associations, can also be prioritized into different functional states depending on their task relevance, and if so how these different formats relate to each other. In the present study, we investigated whether novel WM-guided actions can be brought into different functional states depending on current task demands. Our results reveal that planned actions can be flexibly prioritized when needed and show how their functional state modulates their influence on ongoing behavior. Moreover, they suggest the representations of novel actions of different functional states are maintained in WM via a non-orthogonal coding scheme, thus are prone to interference.

Adaptive coding of stimulus information in human frontoparietal cortex during visual classification.

The neural mechanisms of how frontal and parietal brain regions support flexible adaptation of behavior remain poorly understood. Here, we used functional magnetic resonance imaging (fMRI) and representational similarity analysis (RSA) to investigate frontoparietal representations of stimulus information during visual classification under varying task demands. Based on prior research, we predicted that increasing perceptual task difficulty should lead to adaptive changes in stimulus coding: task-relevant category information should be stronger, while task-irrelevant exemplar-level stimulus information should become weaker, reflecting a focus on the behaviorally relevant category information. Counter to our expectations, however, we found no evidence for adaptive changes in category coding. We did find weakened coding at the exemplar-level within categories however, demonstrating that task-irrelevant information is de-emphasized in frontoparietal cortex. These findings reveal adaptive coding of stimulus information at the exemplar-level, highlighting how frontoparietal regions might support behavior even under challenging conditions.

Theta-phase connectivity between medial prefrontal and posterior areas underlies novel instructions implementation.

Implementing novel instructions is a complex and uniquely human cognitive ability, that requires the rapid and flexible conversion of symbolic content into a format that enables the execution of the instructed behavior. Preparing to implement novel instructions, as opposed to their mere maintenance, involves the activation of the instructed motor plans, and the binding of the action information to the specific context in which this should be executed. Recent evidence and prominent computational models suggest that this efficient configuration of the system might involve a central role of frontal theta oscillations in establishing top-down long-range synchronization between distant and task-relevant brain areas. In the present EEG study (human subjects, 30 females, 4 males), we demonstrate that proactively preparing for the implementation of novels instructions, as opposed to their maintenance, involves a strengthened degree of connectivity in the theta frequency range between medial prefrontal and motor/visual areas. Moreover, we replicated previous results showing oscillatory features associated specifically with implementation demands, and extended on them demonstrating the role of theta oscillations in mediating the effect of task demands on behavioral performance. Taken together, these findings support our hypothesis that the modulation of connectivity patterns between frontal and task-relevant posterior brain areas is a core factor in the emergence of a behavior-guiding format from novel instructions.Significance statementEveryday life requires the use and manipulation of currently available information to guide behavior and reach specific goals. In the present study we investigate how the same instructed content elicits different neural activity depending on the task being performed. Crucially, connectivity between medial prefrontal cortex and posterior brain areas is strengthened when novel instructions have to be implemented, rather than simply maintained. This finding suggests that theta oscillations play a role in setting up a dynamic and flexible network of task-relevant regions optimized for the execution of the instructed behavior.

Canonical template tracking: Measuring the activation state of specific neural representations.

Multivariate analyses of neural data have become increasingly influential in cognitive neuroscience since they allow to address questions about the representational signatures of neurocognitive phenomena. Here, we describe Canonical Template Tracking: a multivariate approach that employs independent localizer tasks to assess the activation state of specific representations during the execution of cognitive paradigms. We illustrate the benefits of this methodology in characterizing the particular content and format of task-induced representations, comparing it with standard (cross-)decoding and representational similarity analyses. Then, we discuss relevant design decisions for experiments using this analysis approach, focusing on the nature of the localizer tasks from which the canonical templates are derived. We further provide a step-by-step tutorial of this method, stressing the relevant analysis choices for functional magnetic resonance imaging and magneto/electroencephalography data. Importantly, we point out the potential pitfalls linked to canonical template tracking implementation and interpretation of the results, together with recommendations to mitigate them. To conclude, we provide some examples from previous literature that highlight the potential of this analysis to address relevant theoretical questions in cognitive neuroscience.

Neural oscillations track the maintenance and proceduralization of novel instructions.

Humans are capable of flexibly converting symbolic instructions into novel behaviors. Previous evidence and theoretical models suggest that the implementation of a novel instruction requires the reformatting of its declarative content into an action-oriented code optimized for the execution of the instructed behavior. While neuroimaging research focused on identifying the brain areas involved in such a process, the temporal and electrophysiological mechanisms remain poorly understood. These mechanisms, however, can provide information about the specific cognitive processes that characterize the proceduralization of information. In the present study, we recorded EEG activity while we asked participants to either simply maintain declaratively the content of novel S-R mappings or to proactively prepare for their implementation. By means of time-frequency analyses, we isolated the oscillatory features specific to the proceduralization of instructions. Implementation of the instructed mappings elicited stronger theta activity over frontal electrodes and suppression in mu and beta activity over central electrodes. On the contrary, activity in the alpha band, which has been shown to track the attentional deployment to task-relevant items, showed no differences between tasks. Together, these results support the idea that proceduralization of information is characterized by specific component processes such as orchestrating complex task settings and configuring the motor system that are not observed when instructions are held in a declarative format.

Frontoparietal action-oriented codes support novel instruction implementation.

A key aspect of human cognitive flexibility concerns the ability to convert complex symbolic instructions into novel behaviors. Previous research proposes that this transformation is supported by two neurocognitive states: an initial declarative maintenance of task knowledge, and an implementation state necessary for optimal task execution. Furthermore, current models predict a crucial role of frontal and parietal brain regions in this process. However, whether declarative and procedural signals independently contribute to implementation remains unknown. We report the results of an fMRI experiment in which participants executed novel instructed stimulus-response associations. We then used a pattern-tracking procedure to quantify the contribution of format-unique signals during instruction implementation. This revealed independent procedural and declarative representations of novel S-Rs in frontoparietal areas, prior to execution. Critically, the degree of procedural activation predicted subsequent behavioral performance. Altogether, our results suggest an important contribution of frontoparietal regions to the neural architecture that regulates cognitive flexibility.

The effects of declaratively maintaining and proactively proceduralizing novel stimulus-response mappings.

Working memory (WM) allows for the maintenance and manipulation of information when carrying out ongoing tasks. Recent models propose that representations in WM can be either in a declarative format (as content of thought) or in a procedural format (in an action-oriented state that drives the cognitive operation to be performed). Current views on the implementation of novel instructions also acknowledge this distinction, assuming these are first encoded as declarative content, and then reformatted into an action-oriented procedural representation upon task demands. Although it is widely accepted that WM has a limited capacity, little is known about the reciprocal costs of maintaining instructions in a declarative format and transforming them in an action code. In a series of three experiments, we asked participants to memorize two or four S-R mappings (i.e., declarative load), and then selected a subset of them by means of a retro-cue to trigger their reformatting into an action-oriented format (i.e., procedural load). We measured the performance in the implementation of the proceduralized mapping and in the declarative recall of the entire set of memorized mappings, to test how the increased load on one component affected the functioning of the other. Our results showed a strong influence of declarative load on the processing of the procedural component, but no effects in the opposite direction. This pattern of results suggests an asymmetry in the costs of maintenance and manipulation in WM, at least when procedural representations cannot be retrieved from long term memory and need to be reformatted online. The available resources seem to be first deployed for the maintenance of all the task-relevant declarative content, and proceduralization takes place to the extent the system can direct attention to the relevant instruction.

Attentional prioritization reconfigures novel instructions into action-oriented task sets.

An astonishing aspect of human cognitive flexibility concerns the ability to efficiently convert complex symbolic instructions into novel behaviors. In such ability, the rapid transformation of relevant content into action plans is particularly crucial as it allows for reflexive, automatic-like execution of merely instructed task sets. However, little is known about the mechanisms that give rise to this transformation. In the current study, we test the hypothesis that novel instructions held in working memory are reformatted into action-oriented representations when selective attention prioritizes their content. To do so, we devised a paradigm in which participants first encoded 4 S-R mappings and later, a retro-cue selected two of them. We first found that participants can benefit from retro-cues during the implementation of novel task-sets. Then, across two preregistered experiments, we observed that cued mappings (but not uncued ones) induced intention-based reflexivity, suggesting that only these entered an action-oriented state. Altogether, our results reveal that selective attention prioritizes relevant novel instructed content, playing an important role in its prospective reformatting into an action-bound task set.

The Neural Bases of Event Monitoring across Domains: a Simultaneous ERP-fMRI Study.

The ability to check and evaluate the environment over time with the aim to detect the occurrence of target stimuli is supported by sustained/tonic as well as transient/phasic control processes, which overall might be referred to as event monitoring. The neural underpinning of sustained attentional control processes involves a fronto-parietal network. However, it has not been well-defined yet whether this cortical circuit acts irrespective of the specific material to be monitored and whether this mediates sustained as well as transient monitoring processes. In the current study, the functional activity of brain during an event monitoring task was investigated and compared between two cognitive domains, whose processing is mediated by differently lateralized areas. Namely, participants were asked to monitor sequences of either faces (supported by right-hemisphere regions) or tools (left-hemisphere). In order to disentangle sustained from transient components of monitoring, a simultaneous EEG-fMRI technique was adopted within a block design. When contrasting monitoring versus control blocks, the conventional fMRI analysis revealed the sustained involvement of bilateral fronto-parietal regions, in both task domains. Event-related potentials (ERPs) showed a more positive amplitude over frontal sites in monitoring compared to control blocks, providing evidence of a transient monitoring component. The joint ERP-fMRI analysis showed that, in the case of face monitoring, this transient component relies on right-lateralized areas, including the inferior parietal lobule and the middle frontal gyrus. In the case of tools, no fronto-parietal areas correlated with the transient ERP activity, suggesting that in this domain phasic monitoring processes were masked by tonic ones. Overall, the present findings highlight the role of bilateral fronto-parietal regions in sustained monitoring, independently of the specific task requirements, and suggest that right-lateralized areas subtend transient monitoring processes, at least in some task contexts.