Levelling up the study of animal gameplay

Play in humans and other animals is widespread and intuitive to recognise. Creative, unstructured play is difficult to quantify, but games direct play towards a specific goal and have defined rules, mechanics and rewards. To date, games have been under-utilised in human and animal behavioural neuroscience. This review evaluates evidence that animals can play human games, including game-theory contests, tangible games, and video games. Animals can be trained to play various human games with cognitive capacities such as role adoption, rule-following and performance monitoring. Animals can make irrational gameplay decisions that jeopardise rewards and have salient emotional responses to winning and losing. Games can advance the field of behavioural neuroscience in several ways. Cognitive tasks can become more engaging and ecologically relevant by adding game elements, known as gamification. Games can be used to induce and measure more naturalistic emotional responses to the process of overcoming (progression/regression) and end state (winning/losing) of cognitive challenges. There is also scope to target specific cognitive skill deficiencies in captive animals using games. However, a recent rapid increase in computerised testing environments raises an important ethical question about the boundary between games and reality for animals.