The free-energy principle (FEP) is a formal model of neuronal processes that is widely recognised in neuroscience as a unifying theory of the brain and biobehaviour. More recently, however, it has been extended beyond the brain to explain the dynamics of living systems, and their unique capacity to avoid decay. The aim of this review is to synthesise these advances with a meta-theoretical ontology of biological systems called variational neuroethology, which integrates the FEP with Tinbergen's four research questions to explain biological systems across spatial and temporal scales. We exemplify this framework by applying it to Homo sapiens, before translating variational neuroethology into a systematic research heuristic that supplies the biological, cognitive, and social sciences with a computationally tractable guide to discovery.

In this paper we outline a framework for the study of the mechanisms involved in the engagement of human agents with cultural affordances. Our aim is to better understand how culture and context interact with human biology to shape human behavior, cognition, and experience. We attempt to integrate several related approaches in the study of the embodied, cognitive, and affective substrates of sociality and culture and the sociocultural scaffolding of experience. The integrative framework we propose bridges cognitive and social sciences to provide (i) an expanded concept of ‘affordance’ that extends to sociocultural forms of life, and (ii) a multilevel account of the socioculturally scaffolded forms of affordance learning and the transmission of affordances in patterned sociocultural practices and regimes of shared attention. This framework provides an account of how cultural content and normative practices are built on a foundation of contentless basic mental processes that acquire content through immersive participation of the agent in social practices that regulate joint attention and shared intentionality.

The processes underwriting the acquisition of culture remain unclear. How are shared habits, norms, and expectations learned and maintained with precision and reliability across large-scale sociocultural ensembles? Is there a unifying account of the mechanisms involved in the acquisition of culture? Notions such as “shared expectations,” the “selective patterning of attention and behaviour,” “cultural evolution,” “cultural inheritance,” and “implicit learning” are the main candidates to underpin a unifying account of cognition and the acquisition of culture; however, their interactions require greater specification and clarification. In this article, we integrate these candidates using the variational (free-energy) approach to human cognition and culture in theoretical neuroscience. We describe the construction by humans of social niches that afford epistemic resources called cultural affordances. We argue that human agents learn the shared habits, norms, and expectations of their culture through immersive participation in patterned cultural practices that selectively pattern attention and behaviour. We call this process “thinking through other minds” (TTOM) – in effect, the process of inferring other agents’ expectations about the world and how to behave in social context. We argue that for humans, information from and about other people's expectations constitutes the primary domain of statistical regularities that humans leverage to predict and organize behaviour. The integrative model we offer has implications that can advance theories of cognition, enculturation, adaptation, and psychopathology. Crucially, this formal (variational) treatment seeks to resolve key debates in current cognitive science, such as the distinction between internalist and externalist accounts of theory of mind abilities and the more fundamental distinction between dynamical and representational accounts of enactivism.

In evolutionary biology, niche construction is sometimes described as a genuine evolutionary process whereby organisms, through their activities and regulatory mechanisms, modify their environment such as to steer their own evolutionary trajectory, and that of other species. There is ongoing debate, however, on the extent to which niche construction ought to be considered a bona fide evolutionary force, on a par with natural selection. Recent formulations of the variational free-energy principle as applied to the life sciences describe the properties of living systems, and their selection in evolution, in terms of variational inference. We argue that niche construction can be described using a variational approach. We propose new arguments to support the niche construction perspective, and to extend the variational approach to niche construction to current perspectives in various scientific fields.

The aim of this article is to clarify how best to interpret some of the central constructs that underwrite the free-energy principle (FEP) -and its corollary, active inference -in theoretical neuroscience and biology: namely, the role that generative models and variational densities play in this theory. We argue that these constructs have been systematically misrepresented in the literature, because of the conflation between the FEP and active inference, on the one hand, and distinct (albeit closely related) Bayesian formulations, centred on the brain -variously known as predictive processing, predictive coding or the prediction error minimisation framework. More specifically, we examine two contrasting interpretations of these models: a structural representationalist interpretation and an enactive interpretation. We argue that the structural representationalist interpretation of generative and recognition models does not do justice to the role that these constructs play in active inference under the FEP. We propose an enactive interpretation of active inferencewhat might be called enactive inference. In active inference under the FEP, the generative and recognition models are best cast as realising inference and control -the self-organising, belief-guided selection of action policies -and do not have the properties ascribed by structural representationalists.

The positive-negative axis of emotional valence has long been recognized as fundamental to adaptive behavior, but its origin and underlying function have largely eluded formal theorizing and computational modeling. Using deep active inference, a hierarchical inference scheme that rests on inverting a model of how sensory data are generated, we develop a principled Bayesian model of emotional valence. This formulation asserts that agents infer their valence state based on the expected precision of their action model—an internal estimate of overall model fitness (“subjective fitness”). This index of subjective fitness can be estimated within any environment and exploits the domain generality of second-order beliefs (beliefs about beliefs). We show how maintaining internal valence representations allows the ensuing affective agent to optimize confidence in action selection preemptively. Valence representations can in turn be optimized by leveraging the (Bayes-optimal) updating term for subjective fitness, which we label affective charge (AC). AC tracks changes in fitness estimates and lends a sign to otherwise unsigned divergences between predictions and outcomes. We simulate the resulting affective inference by subjecting an in silico affective agent to a T-maze paradigm requiring context learning, followed by context reversal. This formulation of affective inference offers a principled account of the link between affect, (mental) action, and implicit metacognition. It characterizes how a deep biological system can infer its affective state and reduce uncertainty about such inferences through internal action (i.e., top-down modulation of priors that underwrite confidence). Thus, we demonstrate the potential of active inference to provide a formal and computationally tractable account of affect. Our demonstration of the face validity and potential utility of this formulation represents the first step within a larger research program. Next, this model can be leveraged to test the hypothesized role of valence by fitting the model to behavioral and neuronal responses.

How do humans come to acquire shared expectations about how they ought to behave in distinct normalized social settings? This paper offers a normative framework to answer this question. We introduce the computational construct of ‘deontic value’ – based on active inference and Markov decision processes – to formalize conceptions of social conformity and human decision-making. Deontic value is an attribute of choices, behaviors, or action sequences that inherit directly from deontic cues in our econiche (e.g., red traffic lights); namely, cues that denote an obligatory social rule. Crucially, the prosocial aspect of deontic value rests upon a particular form of circular causality: deontic cues exist in the environment in virtue of the environment being modified by repeated actions, while action itself is contingent upon the deontic value of environmental cues. We argue that this construction of deontic cues enables the epistemic (i.e., information-seeking) and pragmatic (i.e., goal- seeking) values of any behavior to be ‘cached’ or ‘outsourced’ to the environment, where the environment effectively ‘learns’ about the behavior of its denizens. We describe the process whereby this particular aspect of value enables learning of habitual behavior over neurodevelopmental and transgenerational timescales.

This article presents a unifying theory of the embodied, situated human brain called the Hierarchically Mechanistic Mind (HMM). The HMM describes the brain as a complex adaptive system that actively minimises the decay of our sensory and physical states by producing self-fulfilling action-perception cycles via dynamical interactions between hierarchically organised neurocognitive mechanisms. This theory synthesises the free-energy principle (FEP) in neuroscience with an evolutionary systems theory of psychology that explains our brains, minds, and behaviour by appealing to Tinbergen's four questions: adaptation, phylogeny, ontogeny, and mechanism. After leveraging the FEP to formally define the HMM across different spatiotemporal scales, we conclude by exploring its implications for theorising and research in the sciences of the mind and behaviour.