Modular-integrative modeling: a new framework for building brain models that blend biological realism and functional performance.

This Perspective presents the Modular-Integrative Modeling approach, a novel framework in neuroscience for developing brain models that blend biological realism with functional performance to provide a holistic view on brain function in interaction with the body and environment.

Neural representation in active inference: Using generative models to interact with-and understand-the lived world.

This paper considers neural representation through the lens of active inference, a normative framework for understanding brain function. It delves into how living organisms employ generative models to minimize the discrepancy between predictions and observations (as scored with variational free energy). The ensuing analysis suggests that the brain learns generative models to navigate the world adaptively, not (or not solely) to understand it. Different living organisms may possess an array of generative models, spanning from those that support action-perception cycles to those that underwrite planning and imagination; namely, from explicit models that entail variables for predicting concurrent sensations, like objects, faces, or people-to action-oriented models that predict action outcomes. It then elucidates how generative models and belief dynamics might link to neural representation and the implications of different types of generative models for understanding an agent's cognitive capabilities in relation to its ecological niche. The paper concludes with open questions regarding the evolution of generative models and the development of advanced cognitive abilities-and the gradual transition from pragmatic to detached neural representations. The analysis on offer foregrounds the diverse roles that generative models play in cognitive processes and the evolution of neural representation.

Active inference as a theory of sentient behavior.

This review paper offers an overview of the history and future of active inference-a unifying perspective on action and perception. Active inference is based upon the idea that sentient behavior depends upon our brains' implicit use of internal models to predict, infer, and direct action. Our focus is upon the conceptual roots and development of this theory of (basic) sentience and does not follow a rigid chronological narrative. We trace the evolution from Helmholtzian ideas on unconscious inference, through to a contemporary understanding of action and perception. In doing so, we touch upon related perspectives, the neural underpinnings of active inference, and the opportunities for future development. Key steps in this development include the formulation of predictive coding models and related theories of neuronal message passing, the use of sequential models for planning and policy optimization, and the importance of hierarchical (temporally) deep internal (i.e., generative or world) models. Active inference has been used to account for aspects of anatomy and neurophysiology, to offer theories of psychopathology in terms of aberrant precision control, and to unify extant psychological theories. We anticipate further development in all these areas and note the exciting early work applying active inference beyond neuroscience. This suggests a future not just in biology, but in robotics, machine learning, and artificial intelligence.

Information-theoretical analysis of the neural code for decoupled face representation.

Processing faces accurately and efficiently is a key capability of humans and other animals that engage in sophisticated social tasks. Recent studies reported a decoupled coding for faces in the primate inferotemporal cortex, with two separate neural populations coding for the geometric position of (texture-free) facial landmarks and for the image texture at fixed landmark positions, respectively. Here, we formally assess the efficiency of this decoupled coding by appealing to the information-theoretic notion of description length, which quantifies the amount of information that is saved when encoding novel facial images, with a given precision. We show that despite decoupled coding describes the facial images in terms of two sets of principal components (of landmark shape and image texture), it is more efficient (i.e., yields more information compression) than the encoding in terms of the image principal components only, which corresponds to the widely used eigenface method. The advantage of decoupled coding over eigenface coding increases with image resolution and is especially prominent when coding variants of training set images that only differ in facial expressions. Moreover, we demonstrate that decoupled coding entails better performance in three different tasks: the representation of facial images, the (daydream) sampling of novel facial images, and the recognition of facial identities and gender. In summary, our study provides a first principle perspective on the efficiency and accuracy of the decoupled coding of facial stimuli reported in the primate inferotemporal cortex.

Generating meaning: active inference and the scope and limits of passive AI.

Prominent accounts of sentient behavior depict brains as generative models of organismic interaction with the world, evincing intriguing similarities with current advances in generative artificial intelligence (AI). However, because they contend with the control of purposive, life-sustaining sensorimotor interactions, the generative models of living organisms are inextricably anchored to the body and world. Unlike the passive models learned by generative AI systems, they must capture and control the sensory consequences of action. This allows embodied agents to intervene upon their worlds in ways that constantly put their best models to the test, thus providing a solid bedrock that is - we argue - essential to the development of genuine understanding. We review the resulting implications and consider future directions for generative AI.

Interoceptive technologies for psychiatric interventions: From diagnosis to clinical applications.

Interoception-the perception of internal bodily signals-has emerged as an area of interest due to its implications in emotion and the prevalence of dysfunctional interoceptive processes across psychopathological conditions. Despite the importance of interoception in cognitive neuroscience and psychiatry, its experimental manipulation remains technically challenging. This is due to the invasive nature of existing methods, the limitation of self-report and unimodal measures of interoception, and the absence of standardized approaches across disparate fields. This article integrates diverse research efforts from psychology, physiology, psychiatry, and engineering to address this oversight. Following a general introduction to the neurophysiology of interoception as hierarchical predictive processing, we review the existing paradigms for manipulating interoception (e.g., interoceptive modulation), their underlying mechanisms (e.g., interoceptive conditioning), and clinical applications (e.g., interoceptive exposure). We suggest a classification for interoceptive technologies and discuss their potential for diagnosing and treating mental health disorders. Despite promising results, considerable work is still needed to develop standardized, validated measures of interoceptive function across domains and before these technologies can translate safely and effectively to clinical settings.

Deep kinematic inference affords efficient and scalable control of bodily movements.

Performing goal-directed movements requires mapping goals from extrinsic (workspace-relative) to intrinsic (body-relative) coordinates and then to motor signals. Mainstream approaches based on optimal control realize the mappings by minimizing cost functions, which is computationally demanding. Instead, active inference uses generative models to produce sensory predictions, which allows a cheaper inversion to the motor signals. However, devising generative models to control complex kinematic chains like the human body is challenging. We introduce an active inference architecture that affords a simple but effective mapping from extrinsic to intrinsic coordinates via inference and easily scales up to drive complex kinematic chains. Rich goals can be specified in both intrinsic and extrinsic coordinates using attractive or repulsive forces. The proposed model reproduces sophisticated bodily movements and paves the way for computationally efficient and biologically plausible control of actuated systems.

Information foraging with an oracle.

During ecological decisions, such as when foraging for food or selecting a weekend activity, we often have to balance the costs and benefits of exploiting known options versus exploring novel ones. Here, we ask how individuals address such cost-benefit tradeoffs during tasks in which we can either explore by ourselves or seek external advice from an oracle (e.g., a domain expert or recommendation system). To answer this question, we designed two studies in which participants chose between inquiring (at a cost) for expert advice from an oracle, or to search for options without guidance, under manipulations affecting the optimal choice. We found that participants showed a greater propensity to seek expert advice when it was instrumental to increase payoff (study A), and when it reduced choice uncertainty, above and beyond payoff maximization (study B). This latter result was especially apparent in participants with greater trait-level intolerance of uncertainty. Taken together, these results suggest that we seek expert advice for both economic goals (i.e., payoff maximization) and epistemic goals (i.e., uncertainty minimization) and that our decisions to ask or not ask for advice are sensitive to cost-benefit tradeoffs.

Risky decisions are influenced by individual attributes as a function of risk preference.

It has long been assumed in economic theory that multi-attribute decisions involving several attributes or dimensions - such as probabilities and amounts of money to be earned during risky choices - are resolved by first combining the attributes of each option to form an overall expected value and then comparing the expected values of the alternative options, using a unique evidence accumulation process. A plausible alternative would be performing independent comparisons between the individual attributes and then integrating the results of the comparisons afterwards. Here, we devise a novel method to disambiguate between these types of models, by orthogonally manipulating the expected value of choice options and the relative salience of their attributes. Our results, based on behavioral measures and drift-diffusion models, provide evidence in favor of the framework where information about individual attributes independently impacts deliberation. This suggests that risky decisions are resolved by running in parallel multiple comparisons between the separate attributes - possibly alongside an additional comparison of expected value. This result stands in contrast with the assumption of standard economic theory that choices require a unique comparison of expected values and suggests that at the cognitive level, decision processes might be more distributed than commonly assumed. Beyond our planned analyses, we also discovered that attribute salience affects people of different risk preference type in different ways: risk-averse participants seem to focus more on probability, except when monetary amount is particularly high; risk-neutral/seeking participants, in contrast, seem to focus more on monetary amount, except when probability is particularly low.

Active Vision in Binocular Depth Estimation: A Top-Down Perspective.

Depth estimation is an ill-posed problem; objects of different shapes or dimensions, even if at different distances, may project to the same image on the retina. Our brain uses several cues for depth estimation, including monocular cues such as motion parallax and binocular cues such as diplopia. However, it remains unclear how the computations required for depth estimation are implemented in biologically plausible ways. State-of-the-art approaches to depth estimation based on deep neural networks implicitly describe the brain as a hierarchical feature detector. Instead, in this paper we propose an alternative approach that casts depth estimation as a problem of active inference. We show that depth can be inferred by inverting a hierarchical generative model that simultaneously predicts the eyes' projections from a 2D belief over an object. Model inversion consists of a series of biologically plausible homogeneous transformations based on Predictive Coding principles. Under the plausible assumption of a nonuniform fovea resolution, depth estimation favors an active vision strategy that fixates the object with the eyes, rendering the depth belief more accurate. This strategy is not realized by first fixating on a target and then estimating the depth; instead, it combines the two processes through action-perception cycles, with a similar mechanism of the saccades during object recognition. The proposed approach requires only local (top-down and bottom-up) message passing, which can be implemented in biologically plausible neural circuits.

Goal-seeking compresses neural codes for space in the human hippocampus and orbitofrontal cortex.

Humans can navigate flexibly to meet their goals. Here, we asked how the neural representation of allocentric space is distorted by goal-directed behavior. Participants navigated an agent to two successive goal locations in a grid world environment comprising four interlinked rooms, with a contextual cue indicating the conditional dependence of one goal location on another. Examining the neural geometry by which room and context were encoded in fMRI signals, we found that map-like representations of the environment emerged in both hippocampus and neocortex. Cognitive maps in hippocampus and orbitofrontal cortices were compressed so that locations cued as goals were coded together in neural state space, and these distortions predicted successful learning. This effect was captured by a computational model in which current and prospective locations are jointly encoded in a place code, providing a theory of how goals warp the neural representation of space in macroscopic neural signals.

Beyond simple laboratory studies: Developing sophisticated models to study rich behavior.

Psychology and neuroscience are concerned with the study of behavior, of internal cognitive processes, and their neural foundations. However, most laboratory studies use constrained experimental settings that greatly limit the range of behaviors that can be expressed. While focusing on restricted settings ensures methodological control, it risks impoverishing the object of study: by restricting behavior, we might miss key aspects of cognitive and neural functions. In this article, we argue that psychology and neuroscience should increasingly adopt innovative experimental designs, measurement methods, analysis techniques and sophisticated computational models to probe rich, ecologically valid forms of behavior, including social behavior. We discuss the challenges of studying rich forms of behavior as well as the novel opportunities offered by state-of-the-art methodologies and new sensing technologies, and we highlight the importance of developing sophisticated formal models. We exemplify our arguments by reviewing some recent streams of research in psychology, neuroscience and other fields (e.g., sports analytics, ethology and robotics) that have addressed rich forms of behavior in a model-based manner. We hope that these "success cases" will encourage psychologists and neuroscientists to extend their toolbox of techniques with sophisticated behavioral models - and to use them to study rich forms of behavior as well as the cognitive and neural processes that they engage.

Strategic Task Decomposition in Joint Action.

The core of human cooperation is people's ability to perform joint actions. Frequently, this requires effectively decomposing a joint task into individual subtasks, for example, when jointly shopping at the market to buy food. Surprisingly, little is known about how collaborators balance the costs of establishing a joint strategy for such decompositions and its expected benefits for a joint goal. We created a new online task that required pairs of randomly matched participants to jointly collect colored items. We then systematically varied the cognitive costs and benefits of applying a color-splitting strategy. The results showed that pairs adopted a color-splitting strategy more often when necessary to lower cognitive costs. However, once the strategy was jointly adopted, it continued to be used even when the cost-benefits changed. Our results provide first insights on how people decompose joint tasks into individual components and how decomposition strategies may evolve into conventions.

An active inference model of hierarchical action understanding, learning and imitation.

We advance a novel active inference model of the cognitive processing that underlies the acquisition of a hierarchical action repertoire and its use for observation, understanding and imitation. We illustrate the model in four simulations of a tennis learner who observes a teacher performing tennis shots, forms hierarchical representations of the observed actions, and imitates them. Our simulations show that the agent's oculomotor activity implements an active information sampling strategy that permits inferring the kinematic aspects of the observed movement, which lie at the lowest level of the action hierarchy. In turn, this low-level kinematic inference supports higher-level inferences about deeper aspects of the observed actions: proximal goals and intentions. Finally, the inferred action representations can steer imitative responses, but interfere with the execution of different actions. Our simulations show that hierarchical active inference provides a unified account of action observation, understanding, learning and imitation and helps explain the neurobiological underpinnings of visuomotor cognition, including the multiple routes for action understanding in the dorsal and ventral streams and mirror mechanisms.

The traps of adaptation: Addiction as maladaptive referent-dependent evaluation.

Referent-dependent evaluation theories propose that the ongoing context influences how the brain attributes value to stimuli. What are the implications of these theories for understanding addiction? The paper asks this question by casting this disorder as a form of maladaptive referent-dependent evaluation. Specifically, addiction is proposed to arise from the establishment of an excessive reference point following repeated drug consumption. Several key aspects of the disorder emerge from this perspective, including withdrawal, tolerance, enhanced craving, negative mood, and diminished stimulus discriminability. As highlighted in the paper, this formulation has important analogies with classical accounts of addiction, such as set point theories and associative learning theories. Moreover, this picture fits with the pattern of striatal dopaminergic activity observed in addiction, a key neural signature of the disorder. Overall, the referent-dependent evaluation approach emerges as a useful add-on to the theoretical toolkit adopted to interpret addiction. This also supports the idea that referent-dependent evaluation might offer a general framework to understand various disorders characterised by disrupted motivation.

Cognitive effort and active inference.

This paper aims to integrate some key constructs in the cognitive neuroscience of cognitive control and executive function by formalising the notion of cognitive (or mental) effort in terms of active inference. To do so, we call upon a task used in neuropsychology to assess impulse inhibition-a Stroop task. In this task, participants must suppress the impulse to read a colour word and instead report the colour of the text of the word. The Stroop task is characteristically effortful, and we unpack a theory of mental effort in which, to perform this task accurately, participants must overcome prior beliefs about how they would normally act. However, our interest here is not in overt action, but in covert (mental) action. Mental actions change our beliefs but have no (direct) effect on the outside world-much like deploying covert attention. This account of effort as mental action lets us generate multimodal (choice, reaction time, and electrophysiological) data of the sort we might expect from a human participant engaging in this task. We analyse how parameters determining cognitive effort influence simulated responses and demonstrate that-when provided only with performance data-these parameters can be recovered, provided they are within a certain range.

Evidence or Confidence: What Is Really Monitored during a Decision?

Assessing our confidence in the choices we make is important to making adaptive decisions, and it is thus no surprise that we excel in this ability. However, standard models of decision-making, such as the drift-diffusion model (DDM), treat confidence assessment as a post hoc or parallel process that does not directly influence the choice, which depends only on accumulated evidence. Here, we pursue the alternative hypothesis that what is monitored during a decision is an evolving sense of confidence (that the to-be-selected option is the best) rather than raw evidence. Monitoring confidence has the appealing consequence that the decision threshold corresponds to a desired level of confidence for the choice, and that confidence improvements can be traded off against the resources required to secure them. We show that most previous findings on perceptual and value-based decisions traditionally interpreted from an evidence-accumulation perspective can be explained more parsimoniously from our novel confidence-driven perspective. Furthermore, we show that our novel confidence-driven DDM (cDDM) naturally generalizes to decisions involving any number of alternative options - which is notoriously not the case with traditional DDM or related models. Finally, we discuss future empirical evidence that could be useful in adjudicating between these alternatives.

Humans account for cognitive costs when finding shortcuts: An information-theoretic analysis of navigation.

When faced with navigating back somewhere we have been before we might either retrace our steps or seek a shorter path. Both choices have costs. Here, we ask whether it is possible to characterize formally the choice of navigational plans as a bounded rational process that trades off the quality of the plan (e.g., its length) and the cognitive cost required to find and implement it. We analyze the navigation strategies of two groups of people that are firstly trained to follow a "default policy" taking a route in a virtual maze and then asked to navigate to various known goal destinations, either in the way they want ("Go To Goal") or by taking novel shortcuts ("Take Shortcut"). We address these wayfinding problems using InfoRL: an information-theoretic approach that formalizes the cognitive cost of devising a navigational plan, as the informational cost to deviate from a well-learned route (the "default policy"). In InfoRL, optimality refers to finding the best trade-off between route length and the amount of control information required to find it. We report five main findings. First, the navigational strategies automatically identified by InfoRL correspond closely to different routes (optimal or suboptimal) in the virtual reality map, which were annotated by hand in previous research. Second, people deliberate more in places where the value of investing cognitive resources (i.e., relevant goal information) is greater. Third, compared to the group of people who receive the "Go To Goal" instruction, those who receive the "Take Shortcut" instruction find shorter but less optimal solutions, reflecting the intrinsic difficulty of finding optimal shortcuts. Fourth, those who receive the "Go To Goal" instruction modulate flexibly their cognitive resources, depending on the benefits of finding the shortcut. Finally, we found a surprising amount of variability in the choice of navigational strategies and resource investment across participants. Taken together, these results illustrate the benefits of using InfoRL to address navigational planning problems from a bounded rational perspective.