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.

The study of human–animal interactions has provided insights into the welfare of many species. To date, however, research has largely focused on human relationships with captive mammals, with relatively little exploration of interactions between humans and other vertebrates, despite non-mammals constituting the vast majority of animals currently living under human management. With this study, we aimed to address this gap in knowledge by investigating human–fish interactions at a community garden/aquaponics learning-center that is home to approximately 150 goldfish (Carassius auratus) and seven adult and two juvenile koi (Cyprinus rubrofuscus). After a habituation period (July–September 2019) during which time the fish were regularly provided with the opportunity to engage with the researcher’s submerged hand, but were not forced to interact with the researcher, we collected video data on 10 non-consecutive study days during the month of October. This procedure produced 18~20-min interaction sessions, 10 during T1 (when the experimenter first arrived and the fish had not been fed) and eight during T2 (20–30 min after the fish had been fed to satiation; two sessions of which were lost due equipment malfunction). Interactions between the researcher and the seven adult koi were coded from video based on location (within reach, on the periphery, or out of reach from the researcher) and instances of physical, tactile interaction. Analyses revealed that overall, koi spent more time than expected within reach of the researcher during both T1 (p < 0.02) and T2 (p < 0.03). There were also substantial differences between individuals’ overall propensity for being within-reach and engaging in physical interaction. These results show that koi will voluntarily interact with humans and that individual koi display unique and consistent patterns of interaction. By providing quantitative data to support anecdotal claims that such relationships exist around the world, this research contributes to the ongoing discoveries highlighting the profound dissonance between how humans think about and treat fish and who fish actually are, thereby emphasizing the necessity of stronger moral and legal protections for fishes.