Working memory as an indicator for comparative cognition

Working memory (WM), the representation of information held accessible for manipulation over time, is an essential component of all higher cognitive abilities. It allows for complex behaviors that go beyond simple stimulus-response associations and inflexible behavioral patterns. WM capacity determines how many different pieces of information (items) can be used for these cognitive processes, and in humans, it correlates with fluid intelligence. As such, WM might be a useful tool for comparison of cognition across species. WM can be tested using comparatively simple behavioral protocols, based on operant conditioning, in a multitude of different species. Species-specific contextual variables that influence an animal's performance on a non-cognitive level are controlled by adapting the WM paradigm. The neuronal mechanisms by which WM emerges in the brain, as sustained neuronal activity, are comparable between the different species studied (mammals and birds), as are the areas of the brain in which WM activity can be measured. Thus WM is comparable between vastly different species within their respective niches, accounting for specific contextual variables and unique adaptations. By approaching the question of "general cognitive abilities" or "intelligence" within the animal kingdom from the perspective of WM, the complexity of the core question at hand is reduced to a fundamental memory system required to allow for complex cognitive abilities. This article argues that measuring WM can be a suitable addition to the toolkit of comparative cognition. By measuring WM on a behavioral level and going beyond behavior to the underlying physiological processes, qualitative and quantitative differences in cognition between different animal species can be identified, free of contextual restraints