Modern aquaculture, which produces billions of animals from hundreds of species each year, is generating unprecedented welfare risks.

Traditional animal welfare paradigms have focused on maintaining physical health and mitigating negative impacts to wellbeing. Recently, however, the field has increasingly recognized the importance of positive welfare (i.e., mental and physical states that exceed what is necessary for immediate survival) and accordingly introduced manipulations and indicators of positive welfare for use in agriculture, laboratories, and zoos/aquaria. The creation and monitoring of positive welfare requires an in-depth knowledge of species-specific behavior and biology, which necessitates species-specific or, at a minimum, taxa-specific standards. Research on positive welfare in fish is lagging in this regard and therefore merits further consideration. In this paper, we aim to merge what is already known about positive welfare with the existing fish behavior literature to develop a plan of action for fish welfare research that will ultimately contribute to the development of positive welfare standards and assessment strategies for fish. We begin by exploring the origins of positive welfare research and then outline the physical, psychological and species-specific areas of inquiry that can be investigated in fish. In addition to presenting current findings on fish motivation, emotion, potential sources of positive welfare such as fulfillment of motivational urges (establishing agency, engaging in exploration and learning), and play behavior, we also identify promising areas for future research aimed at developing accurate and appropriate indicators of positive welfare in fish.

We propose that a comparative approach to well-being could be the key to understanding ‘the good life.’ Inspired by current theories of human well-being and animal welfare, we designed a novel test of exploration behavior. Environmentally and socially enriched Long-Evans female rats (N = 60) were trained in four simultaneously presented arms of an eight-arm radial-maze. They learned to expect successes in two arms and failures in the other two. After training, 20 animals remained in enriched housing (enrichment-maintenance) while 40 animals were re-housed in standard, isolated conditions (enrichment-removal). Two weeks later, all animals were re-tested in the maze, initially with access to the four familiar arms only. In the final minute, they also had access to the unfamiliar ambiguous-arms. Though both groups showed significant interest in the ambiguous-arms (P<.0001), the enrichment-maintenance group showed a significantly greater exploratory tendency (P<.01) despite having equivalent levels of activity (P>.3). Thus, we show not only that rats will abandon known rewards and incur risk in order to explore, indicating that exploration is valuable in its own right, but also that individuals with (vs. without) enriched housing conditions are more likely to engage in such exploratory behavior. This novel test contributes to the body of knowledge examining the importance of exploration in humans and other animals; implications for animal welfare and human well-being are discussed.

Laboratory mice(Mus musculus)are typically housed in simple cages consisting of one open space. These standard cages may thwart mouse ability to segregate resting areas from areas where they eliminate, a behaviour that is prevalent across the animal kingdom. No scientific work has directly tested whether mice engage in such segregation behaviour, or whether the ability to do so may have welfare consequences. Here we show that mice, whether housed in standard cages or a complex housing system consisting of three interconnected standard cages, kept nesting and elimination sites highly segregated, with nest and urine co-occurring in the same location only 2% of the time. However, mice in the complex system established these clean and dirty sites in separate cages instead of separate locations within one cage, and carried bedding materials (cellulose pellets) from their nesting cages to their latrine cage. Moreover, mice in the complex system displayed more behaviours associated with positive welfare and were less disturbed by weekly husbandry procedures. We conclude that mice find waste products aversive, and that housing mice in a way that facilitates spatial segregation provides a simple way of allowing the expression of natural behaviours and improving welfare.

As animal personality research becomes more central to the study of animal behavior, there is increasing need for theoretical frameworks addressing its causes and consequences. We propose that regulatory focus theory (RFT) could serve as one such framework while also providing insights into how animal personality relates to welfare. RFT distinguishes between two types of approach motivation: promotion, the motivation to approach gains, and prevention, the motivation to approach or maintain safety. Decades of research have established the utility of RFT as a model of human behavior and recent evidence from zoo-housed primates and laboratory rats has suggested that it may be applicable to nonhuman animal behavior as well. Building on these initial studies, we collected data on 60 rats, Rattus norvegicus, navigating an automated maze that allowed individuals to maintain darkness (indicative of prevention/safety-approach motivation) and/or activate food rewards (indicative of promotion/gain-approach motivation). As predicted, both behaviors showed stable individual differences (Ps <0.01) and were inversely associated with physiological signs of chronic stress, possibly indicating poor welfare (Ps <0.05). Subsequently, half the rats were exposed to a manageable threat (noxious novel object) in the homecage. Re-testing in the maze revealed that threat exposure increased darkness time achieved (P<0.05), suggesting a mechanism by which prevention motivation may be enhanced. These results point toward the potential utility of RFT as a model for animal behavior and welfare.