In this review, we present a new conceptual framework for the study of play behavior, a hitherto puzzling array of seemingly purposeless and unrelated behavioral elements that are recognizable as play throughout the mammalian lineage. Our major new functional hypothesis is that play enables animals to develop flexible kinematic and emotional responses to unexpected events in which they experience a sudden loss of control. Specifically, we propose that play functions to increase the versatility of movements used to recover from sudden shocks such as loss of balance and falling over, and to enhance the ability of animals to cope emotionally with unexpected stressful situations. To obtain this "training for the unexpected," we suggest that animals actively seek and create unexpected situations in play through self-handicapping; that is, deliberately relaxing control over their movements or actively putting themselves into disadvantageous positions and situations. Thus, play is comprised of sequences in which the players switch rapidly between well-controlled movements similar to those used in "serious" behavior and self-handicapping movements that result in temporary loss of control. We propose that this playful switching between in-control and out-of-control elements is cognitively demanding, setting phylogenetic and ontogenetic constraints on play, and is underlain by neuroendocrinological responses that produce a complex emotional state known as "having fun." Furthermore, we propose that play is often prompted by relatively novel or unpredictable stimuli, and is thus related to, although distinct from, exploration. We present 24 predictions that arise from our new theoretical framework, examining the extent to which they are supported by the existing empirical evidence and contrasting them with the predictions of four major alternative hypotheses about play. We argue that our "training for the unexpected" hypothesis can account for some previously puzzling kinematic, structural, motivational, emotional, cognitive, social, ontogenetic, and phylogenetic aspects of play. It may also account for a diversity of individual methods for coping with unexpected misfortunes.

In nature, animals need to actively engage with the environment in order to prosper in survival and reproduction. Hence, agency is a central adaptive characteristic of animal life. In this paper, I propose that from the adaptive/functional point of view, four levels of agency can be distinguished, namely passive/reactive agency (animal being behaviourally passive or purely reactive), action-driven agency (animal behaviourally pursuing current desirable outcomes), competence-building agency (animal engaging with the environment to gain skills and information for future use) and aspirational agency (the animal achieving long-term goals through planning and autobiographical reflection). Recent progress in affective neurobiology indicates that each tier of agency is supported by a different type of affective functioning, at least in the case of mammals. Furthermore, the particular agency levels can be linked to distinct degrees of awareness as defined by recent selfhood theories. Based on this coupling between agency adaptive functioning, affective neurobiology and animal awareness levels, I examine several links between animal agency and animal welfare, including the notion of animal boredom, and discuss how animal agency might be promoted in the restrictive frameworks of intensive animal farming.

The concept of animal welfare is evolving due to progress in our scientific understanding of animal biology and changing societal expectations. Animal welfare science has been primarily concerned with minimizing suffering, but there is growing interest in also promoting positive experiences, grouped under the term positive animal welfare (PAW). However, there are discrepancies in the use of the term PAW. An interdisciplinary group arrived at a consensus that ‘PAW can be defined as the animal flourishing through the experience of predominantly positive mental states and the development of competence and resilience. PAW goes beyond ensuring good physical health and the prevention and alleviation of suffering. It encompasses animals experiencing positive mental states resulting from rewarding experiences, including having choices and opportunities to actively pursue goals and achieve desired outcomes’. The definition also considers individual and species-specific differences. It provides a framework for researchers to investigate PAW and thereby generate innovative, informative and reproducible science. Studies of PAW can contribute to a richer picture of an animal’s life and may elucidate the biological foundations of happiness. The definition creates opportunities to inspire scientific progress in animal biology and to align animal care practices, legislation and markets with societal expectations.