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.

Effects of Experiencing CS-US Pairings on Instructed Fear Reversal.

Fear learning allows us to identify and anticipate aversive events and adapt our behavior accordingly. This is often thought to rely on associative learning mechanisms where an initially neutral conditioned stimulus (CS) is repeatedly paired with an aversive unconditioned stimulus (US), eventually leading to the CS also being perceived as aversive and threatening. Importantly, however, humans also show verbal fear learning. Namely, they have the ability to change their responses to stimuli rapidly through verbal instructions about CS-US pairings. Past research on the link between experience-based and verbal fear learning indicated that verbal instructions about a reversal of CS-US pairings can fully override the effects of previously experienced CS-US pairings, as measured through fear ratings, skin conductance, and fear-potentiated startle. However, it remains an open question whether such instructions can also annul learned CS representations in the brain. Here, we used a fear reversal paradigm (female and male participants) in conjunction with representational similarity analysis of fMRI data to test whether verbal instructions fully override the effects of experienced CS-US pairings in fear-related brain regions or not. Previous research suggests that only the right amygdala should show lingering representations of previously experienced threat ("pavlovian trace"). Unexpectedly, we found evidence for the residual effect of prior CS-US experience to be much more widespread than anticipated, in the amygdala but also cortical regions like the dorsal anterior cingulate or dorsolateral prefrontal cortex. This finding shines a new light on the interaction of different fear learning mechanisms, at times with unexpected consequences.SIGNIFICANCE STATEMENT Humans are able to learn about aversive stimuli both from experience (i.e., repeated pairings of conditioned stimulus (CS) and unconditioned stimulus (US; pavlovian conditioning), and from verbal instructions about stimulus pairings. Understanding how experience-based and verbal learning processes interact is key for understanding the cognitive and neural underpinnings of fear learning. We tested whether prior aversive experiences (CS-US pairings) affected subsequent verbal learning, searching for lingering threat signals after verbal instructions reversed a CS from being threatening to being safe. While past research suggested such threat signals can only be found in the amygdala, we found evidence to be much more widespread, including the medial and lateral PFC. This highlights how experience-based and verbal learning processes interact to support adaptive behavior.

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.

Top-down specific preparatory activations for selective attention and perceptual expectations.

Proactive cognition brain models are mainstream nowadays. Within these, preparation is understood as an endogenous, top-down function that takes place prior to the actual perception of a stimulus and improves subsequent behavior. Neuroimaging has shown the existence of such preparatory activity separately in different cognitive domains, however no research to date has sought to uncover their potential similarities and differences. Two of these, often confounded in the literature, are Selective Attention (information relevance) and Perceptual Expectation (information probability). We used EEG to characterize the mechanisms that pre-activate specific contents in Attention and Expectation. In different blocks, participants were cued to the relevance or to the probability of target categories, faces vs. names, in a gender discrimination task. Multivariate Pattern (MVPA) and Representational Similarity Analyses (RSA) during the preparation window showed that both manipulations led to a significant, ramping-up prediction of the relevant or expected target category. However, classifiers trained with data from one condition did not generalize to the other, indicating the existence of unique anticipatory neural patterns. In addition, a Canonical Template Tracking procedure showed that there was stronger anticipatory perceptual reinstatement for relevance than for expectation blocks. Overall, the results indicate that preparation during attention and expectation acts through distinguishable neural mechanisms. These findings have important implications for current models of brain functioning, as they are a first step towards characterizing and dissociating the neural mechanisms involved in top-down anticipatory processing.

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.

Relating free will beliefs and attitudes.

Most people believe in free will, which is foundational for our sense of agency and responsibility. Past research demonstrated that such beliefs are dynamic, and can be manipulated experimentally. Much less is known about free will attitudes (FWAs; do you value free will?), whether they are equally dynamic, and about their relation to free will beliefs (FWBs). If FWAs were strongly positive, people might be reluctant to revise their beliefs even in the face of strong evidence to do so. In this registered report, we developed a novel measure of FWAs and directly related FWBs and attitudes for the first time. We found FWBs and attitudes to be positively related, although to a lesser degree than determinism or dualism beliefs/attitudes. Nevertheless, an experimental manipulation technique aimed at reducing FWBs (Crick text) showed remarkably specific effects on FWBs only, and no effects on FWAs. Overall, these results provide valuable new insights into laypeople's views on free will by including a novel measure of FWAs. They also provide evidence for the validity of a common experimental technique that has been rightfully criticized in the literature lately.

The effect of task demands on the neural patterns generated by novel instruction encoding.

Verbal instructions allow fast and optimal implementation of novel behaviors. Previous research has shown that different control-related variables structure neural activity in frontoparietal regions during the encoding of novel instructed tasks. However, it is uncertain whether different task goals modulate the organizing effect of these variables. In this study, we investigated whether the neural encoding of three task-relevant variables (dimension integration, response set complexity and target category) is modulated by implementation and memorization demands. To do so, we combined functional Magnetic Resonance Imaging (fMRI), an instruction-following paradigm and multivariate analyses. We addressed how and where distributed activity patterns encoded the instructions' variables and the impact of the implementation and memorization demands on the fidelity of these representations. We further explored the nature of the neural code underpinning this process. Our results reveal, first, that the content of to-be-implemented and to-be-memorized instructions is represented in overlapping brain regions, flexibly using a common neural code across tasks. Importantly, they also suggest that preparing to implement the instructions increases the decodability of task-relevant information in frontoparietal areas, in comparison with memorization demands. Overall, our work emphasizes both similarities and differences in task coding under the two contextual demands. These findings qualify the previous understanding of novel instruction processing, suggesting that representing task attributes in a generalizable code, together with the increase in encoding fidelity induced by the implementation goals, could be key mechanisms for proactive control in novel scenarios.

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.

Exploring the Link between Novel Task Proceduralization and Motor Simulation.

Our ability to generate efficient behavior from novel instructions is critical for our adaptation to changing environments. Despite the absence of previous experience, novel instructed content is quickly encoded into an action-based or procedural format, facilitating automatic task processing. In the current work, we investigated the link between proceduralization and motor simulation, specifically, whether the covert activation of the task-relevant responses is used during the assembly of action-based instructions representations. Across three online experiments, we used a concurrent finger-tapping task to block motor simulation during the encoding of novel stimulus-response (S-R) associations. The overlap between the mappings and the motor task at the response level was manipulated. We predicted a greater impairment at mapping implementation in the overlapping condition, where the mappings' relevant response representations were already loaded by the motor demands, and thus, could not be used in the upcoming task simulation. This hypothesis was robustly supported by the three datasets. Nonetheless, the overlapping effect was not modulated by further manipulations of proceduralization-related variables (preparation demands in Exp.2, mapping novelty in Exp.3). Importantly, a fourth control experiment ruled out that our results were driven by alternative accounts as fatigue or negative priming. Overall, we provided strong evidence towards the involvement of motor simulation during anticipatory task reconfiguration. However, this involvement was rather general, and not restricted to novelty scenarios. Finally, these findings can be also integrated into broader models of anticipatory task control, stressing the role of the motor system during preparation.

Metacognitive Awareness of Difficulty in Action Selection: The Role of the Cingulo-opercular Network.

The question whether and how we are able to monitor our own cognitive states (metacognition) has been a matter of debate for decades. Do we have direct access to our cognitive processes, or can we only infer them indirectly based on their consequences? In the current study, we wanted to investigate the brain circuits that underlie the metacognitive experience of fluency in action selection. To manipulate action-selection fluency, we used a subliminal response priming paradigm. On each trial, both male and female human participants additionally engaged in the metacognitive process of rating how hard they felt it was to respond to the target stimulus. Despite having no conscious awareness of the prime, results showed that participants rated incompatible trials (during which subliminal primes interfered with the required response) to be more difficult than compatible trials (where primes facilitated the required response), reflecting metacognitive awareness of difficulty. This increased sense of subjective difficulty was mirrored by increased activity in the rostral cingulate zone and the anterior insula, two regions that are functionally closely connected. Importantly, this reflected activations that were unique to subjective difficulty ratings and were not explained by RTs or prime-response compatibility. We interpret these findings in light of a possible grounding of the metacognitive judgment of fluency in action selection in interoceptive signals resulting from increased effort.

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.

Induced affective states do not modulate effort avoidance.

Recent research reveals that when faced with alternative lines of action, humans tend to choose the less cognitively demanding one, suggesting that cognitive control is intrinsically registered as costly. This idea is further supported by studies showing that the exertion of cognitive control evokes negative affective states. Despite extensive evidence for mood-induced modulations on control abilities, the impact of affective states on the avoidance of cognitive demand is still unknown. Across two well-powered experiments, we tested the hypothesis that negative affective states would increase the avoidance of cognitively demanding tasks. Contrary to our expectations, induced affective states did not modulate the avoidance of demand, despite having an effect on task performance and subjective experience. Altogether, our results indicate that there are limits to the effect of affective signals on cognitive control and that such interaction might depend on specific affective and control settings.

A Gradient of Sharpening Effects by Perceptual Prior across the Human Cortical Hierarchy.

Prior knowledge profoundly influences perceptual processing. Previous studies have revealed consistent suppression of predicted stimulus information in sensory areas, but how prior knowledge modulates processing higher up in the cortical hierarchy remains poorly understood. In addition, the mechanism leading to suppression of predicted sensory information remains unclear, and studies thus far have revealed a mixed pattern of results in support of either the "sharpening" or "dampening" model. Here, using 7T fMRI in humans (both sexes), we observed that prior knowledge acquired from fast, one-shot perceptual learning sharpens neural representation throughout the ventral visual stream, generating suppressed sensory responses. In contrast, the frontoparietal and default mode networks exhibit similar sharpening of content-specific neural representation, but in the context of unchanged and enhanced activity magnitudes, respectively: a pattern we refer to as "selective enhancement." Together, these results reveal a heretofore unknown macroscopic gradient of prior knowledge's sharpening effect on neural representations across the cortical hierarchy.SIGNIFICANCE STATEMENT A fundamental question in neuroscience is how prior knowledge shapes perceptual processing. Perception is constantly informed by internal priors in the brain acquired from past experiences, but the neural mechanisms underlying this process are poorly understood. To date, research on this question has focused on early visual regions, reporting a consistent downregulation when predicted stimuli are encountered. Here, using a dramatic one-shot perceptual learning paradigm, we observed that prior knowledge results in sharper neural representations across the cortical hierarchy of the human brain through a gradient of mechanisms. In visual regions, neural responses tuned away from internal predictions are suppressed. In frontoparietal regions, neural activity consistent with priors is selectively enhanced. These results deepen our understanding of how prior knowledge informs perception.

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.

Shared Neural Representations of Cognitive Conflict and Negative Affect in the Medial Frontal Cortex.

Influential theories of Medial Frontal Cortex (MFC) function suggest that the MFC registers cognitive conflict as an aversive signal, but no study directly tested this idea. Instead, recent studies suggested that nonoverlapping regions in the MFC process conflict and affect. In this preregistered human fMRI study (male and female), we used MVPAs to identify which regions respond similarly to conflict and aversive signals. The results reveal that, of all conflict- and value-related regions, only the ventral pre-supplementary motor area (or dorsal anterior cingulate cortex) showed a shared neural pattern response to different conflict and affect tasks. These findings challenge recent conclusions that conflict and affect are processed independently, and provide support for integrative views of MFC function.SIGNIFICANCE STATEMENT Multiple theories propose that the MFC, and the dorsal ACC in particular, integrates information related to suboptimal outcomes from different psychological domains (e.g., cognitive control and negative affect) with the aim of adaptively steering behavior. In contrast to recent studies in the field, we provide evidence for the idea that cognitive control and negative affect are integrated in the MFC by showing that a classification algorithm trained on discerning cognitive control (conflict vs no conflict) can predict affect (negative vs positive) in the voxel pattern response of the dorsal ACC/pre-SMA.

Cultural pressure and biased responding in free will attitudes.

Whether you believe free will exists has profound effects on your behaviour, across different levels of processing, from simple motor action to social cognition. It is therefore important to understand which specific lay theories are held in the general public and why. Past research largely focused on investigating free will beliefs (FWB, 'Do you think free will exists?'), but largely ignored a second key aspect: free will attitudes (FWA, 'Do you like/value will?'). Attitudes are often independently predictive of behaviour, relative to beliefs, yet we currently know very little about FWAs in the general public. One key issue is whether such attitudes are subject to biased, socially desirable responding. The vast majority of the general public strongly believes in the existence of free will, which might create cultural pressure to value free will positively as well. In this registered report, we used a very large (N = 1100), open available dataset measuring implicit and explicit attitudes towards free will and determinism to address this issue. Our results indicate that both explicit and implicit attitudes towards free will are more positive than attitudes towards determinism. We also show that people experience cultural pressure to value free will, and to devalue determinism. Yet, we found no strong evidence that this cultural pressure affected either implicit or explicit attitudes in this dataset.

Neural representations of social valence bias economic interpersonal choices.

Prior personal information is highly relevant during social interactions. Such knowledge aids in the prediction of others, and it affects choices even when it is unrelated to actual behaviour. In this investigation, we aimed to study the neural representation of positive and negative personal expectations, how these impact subsequent choices, and the effect of mismatches between expectations and encountered behaviour. We employed functional Magnetic Resonance Imaging in combination with a version of the Ultimatum Game (UG) where participants were provided with information about their partners' moral traits previous to receiving their fair or unfair offers. Univariate and multivariate analyses revealed the implication of the supplementary motor area (SMA) and inferior frontal gyrus (IFG) in the representation of expectations about the partners in the game. Further, these regions also represented the valence of these expectations, together with the ventromedial prefrontal cortex (vmPFC). Importantly, the performance of multivariate classifiers in these clusters correlated with a behavioural choice bias to accept more offers following positive descriptions, highlighting the impact of the valence of the expectations on participants' economic decisions. Altogether, our results suggest that expectations based on social information guide future interpersonal decisions and that the neural representation of such expectations in the vmPFC is related to their influence on behaviour.

Investigating the effect of trustworthiness on instruction-based reflexivity.

Unlike other species, humans are capable of rapidly learning new behavior from a single instruction. While previous research focused on the cognitive processes underlying the rapid, automatic implementation of instructions, the fundamentally social nature of instruction following has remained largely unexplored. Here, we investigated whether instructor trustworthiness modulates instruction implementation using both explicit and reflexive measures. In a first preregistered study, we validated a new paradigm to manipulate the perceived trustworthiness of two different virtual characters and showed that such a manipulation reliably induced implicit associations between the virtual characters and trustworthiness attributes. Moreover, we show that trustworthy instructors are followed more frequently and faster. In two additional preregistered experiments, we tested if trustworthiness towards the instructor influenced the cognitive processes underlying instruction implementation. While we show that verbally conveyed instructions led to automatic instruction implementation, this effect was not modulated by the trustworthiness of the instructor. Thus, we succeeded to design and validate a novel trustworthiness manipulation (Experiment 1) and to create a social variant of the instruction-based reflexivity paradigm (Experiments 2 and 3). However, this instruction-based reflexivity effect was not modulated by the instructors' trustworthiness.

Are all behavioral reward benefits created equally? An EEG-fMRI study.

Reward consistently boosts performance in cognitive tasks. Although many different reward manipulations exist, systematic comparisons are lacking. Reward effects on cognitive control are usually studied using monetary incentive delay (MID; cue-related reward information) or stimulus-reward association (SRA; target-related reward information) tasks. While for MID tasks, evidence clearly implicates reward-triggered global increases in proactive control, it is unclear how reward effects arise in SRA tasks, and in how far such mechanisms overlap during task preparation and target processing. Here, we address these questions with simultaneous EEG-fMRI using a Stroop task with four different block types. In addition to MID and SRA blocks, we used an SRA-task modification with reward-irrelevant cues (C-SRA) and regular reward-neutral Stroop-task blocks. Behaviorally, we observed superior performance for all reward conditions compared to Neutral, and more pronounced reward effects in the SRA and C-SRA blocks, compared to MID blocks. The fMRI data showed similar reward effects in value-related areas for events that signaled reward availability (MID cues and (C-)SRA targets), and comparable reward modulations in cognitive-control regions for all targets regardless of block type. This result pattern was echoed by the EEG data, showing clear markers of valuation and cognitive control, which only differed during task preparation, whereas reward-related modulations during target processing were again comparable across block types. Yet, considering only cue-related fMRI data, C-SRA cues triggered preparatory control processes beyond reward-unrelated MID cues, without simultaneous modulations in typical reward areas, implicating enhanced task preparation that is not directly driven by a concurrent neural reward-anticipation response.

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.