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.

The authors hypothesize that depressed states evolved to minimize risk in social interactions in which individuals perceive that the ratio of their social value to others, and their social burden on others, is at a critically low level. When this ratio reaches a point where social value and social burden are approaching equivalence, the individual is in danger of exclusion from social contexts that, over the course of evolution, have been critical to fitness. Many features of depressed states can be understood in relation to mechanisms that reduce social risk in such circumstances, including (a) hyper-sensitivity to signals of social threat from others, (b) sending signals to others that reduce social risks, and (c) inhibiting risk-seeking (e.g., confident, acquisitive) behaviors. These features are discussed in terms of psychosocial and neurobiological research on depressive phenomena.

With different factors identified for psychological distress and resilience, these findings may help inform the development of tailored mental health interventions and resilience-building programs for this vulnerable population.