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.

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Two important questions in bioacoustics are whether vocal repertoires of animals are graded or discrete and how the vocal expressions are linked to the context of emission. Here we address these questions in an ungulate species. The vocal repertoire of young domestic pigs, Sus scrofa, was quantitatively described based on 1513 calls recorded in 11 situations. We described the acoustic quality of calls with 8 acoustic parameters. Based on these parameters, the k-means clustering method showed a possibility to distinguish either two or five clusters although the call types are rather blurred than strictly discrete. The division of the vocal repertoire of piglets into two call types has previously been used in many experimental studies into pig acoustic communication and the five call types correspond well to previously published partial repertoires in specific situations. Clear links exist between the type of situation, its putative valence, and the vocal expression in that situation. These links can be described adequately both with a set of quantitative acoustic variables and through categorisation into call types. The information about the situation of emission of the calls is encoded through five call types almost as accurately as through the full quantitative description.

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.

In 2012, the World Organisation for Animal Health adopted 10 'General Principles for the Welfare of Animals in Livestock Production Systems' to guide the development of animal welfare standards. The General Principles draw on half a century of scientific research relevant to animal welfare: (1) how genetic selection affects animal health, behaviour and temperament; (2) how the environment influences injuries and the transmission of diseases and parasites; (3) how the environment affects resting, movement and the performance of natural behaviour; (4) the management of groups to minimize conflict and allow positive social contact; (5) the effects of air quality, temperature and humidity on animal health and comfort; (6) ensuring access to feed and water suited to the animals' needs and adaptations; (7) prevention and control of diseases and parasites, with humane euthanasia if treatment is not feasible or recovery is unlikely; (8) prevention and management of pain; (9) creation of positive human-animal relationships; and (10) ensuring adequate skill and knowledge among animal handlers. Research directed at animal welfare, drawing on animal behaviour, stress physiology, veterinary epidemiology and other fields, complements more established fields of animal and veterinary science and helps to create a more comprehensive scientific basis for animal care and management.