Working memory enables us to temporarily store and manipulate information, a crucial function for problem-solving. However, most working memory models emphasize cortical interactions ignoring contributions from subcortical and cerebellar regions. Given the dense connectivity between the cerebellum, subcortex, and cortex, we hypothesize that these regions provide unique contributions during working memory manipulation. We tested this hypothesis using functional Magnetic Resonance Imaging (fMRI) to measure blood oxygen-level dependent (BOLD) activity during a mental rotation task, where participants judged whether rotated pairs of three-dimensional stimuli were identical. Our results revealed a distributed network spanning the cortex, subcortex, and cerebellum that differentiates rotated from non-rotated stimuli and correct from incorrect responses. BOLD recruitment in these regions increased with larger angles of rotation. We observed delayed responses in premotor, subcortical, and cerebellar regions during incorrect trials. These findings suggest that cerebellar and subcortical regions support working memory manipulation, highlighting a broader mechanism by which distributed brain regions interact to coordinate higher cognitive functions. ### Competing Interest Statement The authors have declared no competing interest.

Working memory is critical for manipulating and temporarily storing information during cognitive tasks such as problem-solving. Most models focus primarily on cortical-cortical interactions, neglecting subcortical and cerebellar contributions. Given the extensive connectivity between the cerebellum, subcortex, and cortex, we hypothesize that they contribute distinct, yet complementary, functions during working memory manipulation. To test this, we used functional Magnetic Resonance Imaging (fMRI) to measure blood oxygen-level dependent (BOLD) activity while participants performed a mental rotation task. Our results revealed a distributed network spanning the cortex, subcortex, and cerebellum that differentiates rotated from non-rotated stimuli and correct from incorrect responses. Notably, delayed responses in premotor, subcortical, and cerebellar regions during incorrect trials, suggest that their precise recruitment is crucial for successful working memory manipulation. These findings expand current models of working memory manipulation, revealing the collaborative role of subcortical and cerebellar regions in coordinating higher cognitive functions.

Perceptual updating has been proposed to rely upon evolving activity within a recurrent, distributed thalamocortical network whose interconnections are modulated by bursts of ascending neuromodulatory neurotransmitters, such as noradrenaline. To test this hypothesis mechanistically, we leveraged a combination of pupillometry, fMRI and recurrent neural network modelling of an ambiguous figures task. Qualitative shifts in the perceptual interpretation of an ambiguous image were associated with peaks in pupil diameter, an indirect readout of phasic bursts in neuromodulatory tone. We hypothesized that increases in neuromodulatory tone led to neural gain alteration so as to causally mediate perceptual switches. To test this hypothesis, we trained a recurrent neural network to perform an analogous perceptual categorisation task, and then manipulated the gain of the RNN to mimic the effect of neuromodulatory tone. As predicted, we observed an earlier perceptual shift as a function of heightened gain. Leveraging a low-dimensional readout of the RNN dynamics, we developed two novel predictions: perceptual switches should co-occur with peaks in low-dimensional brain state velocity and with flattened energy landscape dynamics. We used dimensionality-reduced summaries of whole-brain fMRI dynamics to independently confirm each of these predictions. These results support the role of the neuromodulatory system in the large-scale network reconfigurations that mediate abrupt changes in perception.

Contemporary models of perceptual awareness lack tractable neurobiological constraints. Inspired by recent cellular recordings in a mouse model of tactile threshold detection, we constructed a biophysical model of perceptual awareness that incorporated essential features of thalamocortical anatomy and cellular physiology. Our model reproduced, and mechanistically explains, the key in vivo neural and behavioural signatures of perceptual awareness in the mouse model, as well as the response to a set of causal perturbations. We generalised the same model (with identical parameters) to a more complex task - visual rivalry - and found that the same thalamic-mediated mechanism of perceptual awareness determined perceptual dominance. This led to the generation of a set of novel, and directly testable, electrophysiological predictions. Analyses of the model based on dynamical systems theory show that perceptual awareness in simulations of both threshold detection and visual rivalry arises from the emergent systems-level dynamics of thalamocortical loops.

The optimism bias is a cognitive bias where individuals overestimate the likelihood of good outcomes and underestimate the likelihood of bad outcomes. Associated with improved quality of life, optimism bias is considered to be adaptive and is a promising avenue of research for mental health interventions in conditions where individuals lack optimism such as major depressive disorder. Here we lay the groundwork for future research on optimism as an intervention by introducing a domain general formal model of optimism bias, which can be applied in different task settings. Employing the active inference framework, we propose a model of the optimism bias as high precision likelihood biased towards positive outcomes. First, we simulate how optimism may be lost during development by exposure to negative events. We then ground our model in the empirical literature by showing how the developmentally acquired differences in optimism are expressed in a belief updating task typically used to assess optimism bias. Finally, we show how optimism affects action in a modified two-armed bandit task. Our model and the simulations it affords provide a computational basis for understanding how optimism bias may emerge, how it may be expressed in standard tasks used to assess optimism, and how it affects agents' decision-making and actions; in combination, this provides a basis for future research on optimism as a mental health intervention.

There is substantial evidence that neuromodulatory systems critically influence brain state dynamics; however, most work has been purely descriptive. Here, we quantify, using data combining local inactivation of the basal forebrain with simultaneous measurement of resting-state fMRI activity in the macaque, the causal role of long-range cholinergic input to the stabilization of brain states in the cerebral cortex. Local inactivation of the nucleus basalis of Meynert (nbM) leads to a decrease in the energy barriers required for an fMRI state transition in cortical ongoing activity. Moreover, the inactivation of particular nbM sub-regions predominantly affects information transfer in cortical regions known to receive direct anatomical projections. We demonstrate these results in a simple neurodynamical model of cholinergic impact on neuronal firing rates and slow hyperpolarizing adaptation currents. We conclude that the cholinergic system plays a critical role in stabilizing macroscale brain state dynamics.

Consciousness can be conceptualized as varying along at least two dimensions: the global state of consciousness and the content of conscious experience. Here, we highlight the cellular and systems-level contributions of the thalamus to conscious state and then argue for thalamic contributions to conscious content, including the integrated, segregated, and continuous nature of our experience. We underscore vital, yet distinct roles for core- and matrix-type thalamic neurons. Through reciprocal interactions with deep-layer cortical neurons, matrix neurons support wakefulness and determine perceptual thresholds, whereas the cortical interactions of core neurons maintain content and enable perceptual constancy. We further propose that conscious integration, segregation, and continuity depend on the convergent nature of corticothalamic projections enabling dimensionality reduction, a thalamic reticular nucleus-mediated divisive normalization-like process, and sustained coherent activity in thalamocortical loops, respectively. Overall, we conclude that the thalamus plays a central topological role in brain structures controlling conscious experience.

Understanding the neurobiological mechanisms underlying consciousness remains a significant challenge. Recent evidence suggests that the coupling between distal-apical and basal-somatic dendrites in thick-tufted layer 5 pyramidal neurons (L5PN), regulated by the nonspecific-projecting thalamus, is crucial for consciousness. Yet, it is uncertain whether this thalamocortical mechanism can support emergent signatures of consciousness, such as integrated information. To address this question, we constructed a biophysical network of dual-compartment thick-tufted L5PN, with dendrosomatic coupling controlled by thalamic inputs. Our findings demonstrate that integrated information is maximized when nonspecific thalamic inputs drive the system into a regime of time-varying synchronous bursting. Here, the system exhibits variable spiking dynamics with broad pairwise correlations, supporting the enhanced integrated information. Further, the observed peak in integrated information aligns with criticality signatures and empirically observed layer 5 pyramidal bursting rates. These results suggest that the thalamocortical core of the mammalian brain may be evolutionarily configured to optimize effective information processing, providing a potential neuronal mechanism that integrates microscale theories with macroscale signatures of consciousness.

The active inference framework, and in particular its recent formulation as a partially observable Markov decision process (POMDP), has gained increasing popularity in recent years as a useful approach for modeling neurocognitive processes. This framework is highly general and flexible in its ability to be customized to model any cognitive process, as well as simulate predicted neuronal responses based on its accompanying neural process theory. It also affords both simulation experiments for proof of principle and behavioral modeling for empirical studies. However, there are limited resources that explain how to build and run these models in practice, which limits their widespread use. Most introductions assume a technical background in programming, mathematics, and machine learning. In this paper we offer a step-by-step tutorial on how to build POMDPs, run simulations using standard MATLAB routines, and fit these models to empirical data. We assume a minimal background in programming and mathematics, thoroughly explain all equations, and provide exemplar scripts that can be customized for both theoretical and empirical studies. Our goal is to provide the reader with the requisite background knowledge and practical tools to apply active inference to their own research. We also provide optional technical sections and multiple appendices, which offer the interested reader additional technical details. This tutorial should provide the reader with all the tools necessary to use these models and to follow emerging advances in active inference research.

Perception is thought to rely upon evolving activity within a recurrent, distributed thalamocortical network whose interconnections are modulated by bursts of noradrenaline. To test this hypothesis, we leveraged a combination of pupillometry, fMRI and recurrent neural network modelling of an ambiguous figures task. Shifts in the perceptual interpretation of an ambiguous image were associated with peaks in pupil diameter, implicating noradrenergic gain alteration in the perceptual switch. We trained a recurrent neural network to perform an analogous perceptual categorisation task, and then manipulated the gain of the RNN to mimic the effect of noradrenaline. We observed an earlier perceptual shift as a function of heightened gain. Leveraging a low-dimensional readout of the RNN dynamics, and a measure of the energy landscape traversed by the dynamics, we developed two novel predictions: perceptual switches should co-occur with peaks in low-dimensional brain state velocity and with flattened energy landscape dynamics. We used whole-brain fMRI data to confirm these predictions. These results confirm the core role of the noradrenergic system in the large-scale network reconfigurations that mediate perception.

OPINION article Front. Psychol., 18 November 2022Sec. Theoretical and Philosophical Psychology https://doi.org/10.3389/fpsyg.2022.1043747

Cognitive theories of consciousness, such as global workspace theory and higher-order theories, posit that frontoparietal circuits play a crucial role in conscious access. However, recent studies using no-report paradigms have posed a challenge to cognitive theories by demonstrating conscious accessibility in the apparent absence of prefrontal cortex (PFC) activation. To address this challenge, this paper presents a computational model of conscious access, based upon active inference, that treats working memory gating as a cognitive action. We simulate a visual masking task and show that late P3b-like event-related potentials (ERPs), and increased PFC activity, are induced by the working memory demands of report. When reporting demands are removed, these late ERPs vanish and PFC activity is reduced. These results therefore reproduce, and potentially explain, results from no-report paradigms. However, even without reporting demands, our model shows that simulated PFC activity on visible stimulus trials still crosses the threshold for reportability – maintaining the link between PFC and conscious access. Therefore, our simulations show that evidence provided by no-report paradigms does not necessarily contradict cognitive theories of consciousness.

Measuring cognition in single subjects presents unique challenges. On the other hand, individually sensitive measurements offer extraordinary opportunities, from informing theoretical models to enabling truly individualised clinical assessment. Here, we test the robustness of fast, periodic, and visual stimulation (FPVS), an emerging method proposed to elicit detectable responses to written words in the electroencephalogram (EEG) of individual subjects. The method is non-invasive, passive, and requires only a few minutes of testing, making it a potentially powerful tool to test comprehension in those who do not speak or who struggle with long testing procedures. In an initial study, Lochy et al. (2015) used FPVS to detect word processing in eight out of 10 fluent French readers. Here, we attempted to replicate their study in a new sample of 10 fluent English readers. Participants viewed rapid streams of pseudo-words with words embedded at regular intervals, while we recorded their EEG. Based on Lochy et al. (2015) we expected that words would elicit a steady-state response at the word-presentation frequency (2 Hz) over parieto-occipital electrode sites. However, across 40 datasets (10 participants, two conditions, and two regions of interest–ROIs), only four datasets met the criteria for a unique response to words. This corresponds to a 10% detection rate. We conclude that FPVS should be developed further before it can serve as an individually-sensitive measure of written word processing.

Classic models of predictive coding propose that sensory systems use information retained from prior experience to predict current sensory input. Any mismatch between predicted and current input (prediction error) is then fed forward up the hierarchy leading to a revision of the prediction. We tested this hypothesis in the domain of object vision using a combination of multivariate pattern analysis and time-resolved electroencephalography. We presented participants with sequences of images that stepped around fixation in a predictable order. On the majority of presentations, the images conformed to a consistent pattern of position order and object category order, however, on a subset of presentations the last image in the sequence violated the established pattern by either violating the predicted category or position of the object. Contrary to classic predictive coding when decoding position and category we found no differences in decoding accuracy between predictable and violation conditions. However, consistent with recent extensions of predictive coding, exploratory analyses showed that a greater proportion of predictions was made to the forthcoming position in the sequence than to either the previous position or the position behind the previous position suggesting that the visual system actively anticipates future input as opposed to just inferring current input.

The global neuronal workspace (GNW) model has inspired over two decades of hypothesis-driven research on the neural basis of consciousness. However, recent studies have reported findings that are at odds with empirical predictions of the model. Further, the macro-anatomical focus of current GNW research has limited the specificity of predictions afforded by the model. In this paper we present a neurocomputational model - based on Active Inference - that captures central architectural elements of the GNW and is able to address these limitations. The resulting 'predictive global workspace' casts neuronal dynamics as approximating Bayesian inference, allowing precise, testable predictions at both the behavioural and neural levels of description. We report simulations demonstrating the model's ability to reproduce: 1) the electrophysiological and behavioural results observed in previous studies of inattentional blindness; and 2) the previously introduced four-way taxonomy predicted by the GNW, which describes the relationship between consciousness, attention, and sensory signal strength. We then illustrate how our model can reconcile/explain (apparently) conflicting findings, extend the GNW taxonomy to include the influence of prior expectations, and inspire novel paradigms to test associated behavioural and neural predictions.

Classic models of predictive coding propose that sensory systems use information retained from prior experience to predict current sensory input. Any mismatch between predicted and current input (prediction error) is then fed forward up the hierarchy leading to a revision of the prediction. We tested this hypothesis in the domain of object vision using a combination of multivariate pattern analysis and time-resolved electroencephalography. We presented participants with sequences of images that stepped around fixation in a predictable order. On the majority of presentations, the images conformed to a consistent pattern of position order and object category order, however, on a subset of presentations the last image in the sequence violated the established pattern by either violating the predicted category or position of the object. Contrary to classic predictive coding when decoding position and category we found no differences in decoding accuracy between predictable and violation conditions. However, consistent with recent extensions of predictive coding, exploratory analyses showed that a greater proportion of predictions was made to the forthcoming position in the sequence than to either the previous position or the position behind the previous position suggesting that the visual system actively anticipates future input as opposed to just inferring current input.