Concurrent response and action effect representations across the somatomotor cortices during novel task preparation.

Instructions allow us to fulfill novel and complex tasks on the first try. This skill has been linked to preparatory brain signals that encode upcoming demands in advance, facilitating novel performance. To deepen insight into these processes, we explored whether instructions pre-activated task-relevant motoric and perceptual neural states. Critically, we addressed whether these representations anticipated activity patterns guiding overt sensorimotor processing, which could reflect that internally simulating novel tasks facilitates the preparation. To do so, we collected functional magnetic resonance imaging data while female and male participants encoded and implemented novel stimulus-response associations. Participants also completed localizer tasks designed to isolate the neural representations of the mappings-relevant motor responses, perceptual consequences, and stimulus categories. Using canonical template tracking, we identified whether and where these sensorimotor representations were pre-activated. We found that response-related templates were encoded in advance in regions linked with action control, entailing not only the instructed responses but also their somatosensory consequences. This result was particularly robust in primary motor and somatosensory cortices. While, following our predictions, we found a systematic decrease in the irrelevant stimulus templates' representational strength compared to the relevant ones, this difference was due to below-zero estimates linked to the irrelevant category activity patterns. Overall, our findings reflect that instruction processing relies on the sensorimotor cortices to anticipate motoric and kinesthetic representations of prospective action plans, suggesting the engagement of motor imagery during novel task preparation. More generally, they stress that the somatomotor system could participate with higher-level frontoparietal regions during anticipatory task control.

Proactive selective attention across competition contexts.

Selective attention is a cognitive function that helps filter out unwanted information. Theories such as the biased competition model (Desimone & Duncan, 1995) explain how attentional templates bias processing towards targets in contexts where multiple stimuli compete for resources. However, it is unclear how the anticipation of different levels of competition influences the nature of attentional templates, in a proactive fashion. In this study, we used electroencephalography (EEG) to investigate how the anticipated demands of attentional selection (either high or low stimuli competition contexts) modulate target-specific preparatory brain activity and its relationship with task performance. To do so, participants performed a sex/gender judgment task in a cue-target paradigm where, depending on the block, target and distractor stimuli appeared simultaneously (high competition) or sequentially (low competition). Multivariate Pattern Analysis (MVPA) showed that, in both competition contexts, there was a preactivation of the target category to select, with a ramping-up profile at the end of the preparatory interval. However, cross-classification showed no generalization across competition conditions, suggesting different preparatory formats. Notably, time-frequency analyses showed differences between anticipated competition demands, with higher theta band power for high than low competition, which mediated the impact of subsequent stimuli competition on behavioral performance. Overall, our results show that, whereas preactivation of the internal templates associated with the category to select are engaged in advance in high and low competition contexts, their underlying neural patterns differ. In addition, these codes could not be associated with theta power, suggesting that they reflect different preparatory processes. The implications of these findings are crucial to increase our understanding of the nature of top-down processes across different contexts.

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.

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.

The Representations of Chinese Characters: Evidence from Sublexical Components.

Little research has been done about the neural substrate of the sublexical level of Chinese word recognition. In particular, it is unclear how radicals participate in Chinese word processing. We compared two measures of radical combinability, position-general radical combinability (GRC) and position-specific radical combinability (SRC) depending on whether the position of the radical is taken into account. We selected characters with embedded target radicals that had different GRC and SRC measures. These measures were used as predictors in a parametric modulation analysis and a multivariate representational similarity analysis. Human participants with native Mandarin speakers (17 males and 24 females) were asked to read words in search of animal words. Results showed that SRC is a better predictor than GRC in decoding the neural patterns. Whole-brain analysis indicated that SRC is encoded bilaterally in the inferior frontal gyrus (IFG, pars opercularis, and pars triangularis), the middle frontal gyrus (MFG), and a region on the border of the superior parietal lobule and the inferior parietal lobule (SPL/IPL). Region-of-interest-based RSA confirmed the results of the whole-brain analysis. Furthermore, we observed a correlation of another sublexical variable, logographeme composition, with bilateral activity in SPL. Logographemes refer to the basic stroke combinations that form radicals and characters. Finally, we observed involvement of bilateral cerebellum activity in Chinese word recognition. Our findings confirm the importance of sublexical components (SRC and logographeme composition) in Chinese word recognition and also confirm that Chinese word recognition involves more bilateral processing than word recognition in alphabetical languages.SIGNIFICANCE STATEMENT Chinese is a logographic language. However, characters contain informative subword components (radicals, logographemes, and strokes). We investigated whether the position of the radical is important. We presented carefully selected words and looked where brain activity correlated with subword information. Results indicate that position-dependent radicals predict brain encoding in a network of regions associated with Chinese word recognition, including higher order regions such as bilateral IFG, MFG, and SPL/IPL. Logographeme composition had an effect as well. Our findings provide strong evidence (1) for the importance of position-specific radical information and logographemes in Chinese word recognition, (2) that current brain imaging techniques are best suited to study these, and (3) that confirms the interactive nature of Chinese character recognition.

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.

Representational Organization of Novel Task Sets during Proactive Encoding.

Recent multivariate analyses of brain data have boosted our understanding of the organizational principles that shape neural coding. However, most of this progress has focused on perceptual visual regions (Connolly et al., 2012), whereas far less is known about the organization of more abstract, action-oriented representations. In this study, we focused on humans' remarkable ability to turn novel instructions into actions. While previous research shows that instruction encoding is tightly linked to proactive activations in frontoparietal brain regions, little is known about the structure that orchestrates such anticipatory representation. We collected fMRI data while participants (both males and females) followed novel complex verbal rules that varied across control-related variables (integrating within/across stimuli dimensions, response complexity, target category) and reward expectations. Using representational similarity analysis (Kriegeskorte et al., 2008), we explored where in the brain these variables explained the organization of novel task encoding, and whether motivation modulated these representational spaces. Instruction representations in the lateral PFC were structured by the three control-related variables, whereas intraparietal sulcus encoded response complexity and the fusiform gyrus and precuneus organized its activity according to the relevant stimulus category. Reward exerted a general effect, increasing the representational similarity among different instructions, which was robustly correlated with behavioral improvements. Overall, our results highlight the flexibility of proactive task encoding, governed by distinct representational organizations in specific brain regions. They also stress the variability of motivation-control interactions, which appear to be highly dependent on task attributes, such as complexity or novelty.SIGNIFICANCE STATEMENT In comparison with other primates, humans display a remarkable success in novel task contexts thanks to our ability to transform instructions into effective actions. This skill is associated with proactive task-set reconfigurations in frontoparietal cortices. It remains yet unknown, however, how the brain encodes in anticipation the flexible, rich repertoire of novel tasks that we can achieve. Here we explored cognitive control and motivation-related variables that might orchestrate the representational space for novel instructions. Our results showed that different dimensions become relevant for task prospective encoding, depending on the brain region, and that the lateral PFC simultaneously organized task representations following different control-related variables. Motivation exerted a general modulation upon this process, diminishing rather than increasing distances among instruction representations.

Transient and Sustained Control Mechanisms Supporting Novel Instructed Behavior.

The success of humans in novel environments is partially supported by our ability to implement new task procedures via instructions. This complex skill has been associated with the activity of control-related brain areas. Current models link fronto-parietal and cingulo-opercular networks with transient and sustained modes of cognitive control, based on observations during repetitive task settings or rest. The current study extends this dual model to novel instructed tasks. We employed a mixed design and an instruction-following task to extract phasic and tonic brain signals associated with the encoding and implementation of novel verbal rules. We also performed a representation similarity analysis to capture consistency in task-set encoding within trial epochs. Our findings show that both networks are involved while following novel instructions: transiently, during the implementation of the instruction, and in a sustained fashion, across novel trials blocks. Moreover, the multivariate results showed that task representations in the cingulo-opercular network were more stable than in the fronto-parietal one. Our data extend the dual model of cognitive control to novel demanding situations, highlighting the high flexibility of control-related regions in adopting different temporal profiles.

Encoding, preparation and implementation of novel complex verbal instructions.

Verbal instructions allow humans to acquire and implement complex novel rules in few seconds. A major question that remains elusive is how the brain represents this information prior to successful task execution. In this experiment, we studied the brain regions involved in representing categorical stimulus information during the encoding of novel instructions, their preparation and also their implementation, as well as the relation of the fidelity of these representations to observable behavior. To do so, we devised a novel instructions paradigm to delimitate these three stages. Using univariate and multivariate analyses of functional magnetic resonance data, our study revealed that the semantic content (faces or letters) of complex novel instructions can be decoded several seconds before the onset of a target, as soon as instructions are encoded. Crucially, the quality of the information represented in domain-general and category-selective regions correlated with subsequent behavioral performance. This suggests that the rapid transformation of novel instructions into coherent behavior is supported by control mechanisms that use available, relevant information about the current rule prior to its execution. In addition, our results highlight the relation between these control processes and others such as prospective memory and maintenance of future intentions.