People retrieve episodic memories about specific earlier events that happened to them. Accordingly, researchers have sought to evaluate the hypothesis that nonhumans retrieve episodic memories. The central hypothesis of an animal model of episodic memory is that, at the moment of a memory assessment, the animal retrieves a memory of the specific earlier event. Testing this hypothesis requires the elimination of nonepisodic memory hypotheses. A number of case studies focus on the development of animal models of episodic memory, including what-where-when memory, source memory, item-in-context memory, and unexpected questions. Compelling evidence for episodic memory comes from studies in which judgments of familiarity cannot produce accurate choices in memory assessments. These approaches may be used to explore the evolution of cognition.
Keywords: episodic memory, what-where-when memory, source memory, binding, item-in-context memory
IntroductionFundamental aspects of human cognition raise a natural question, namely, How widely distributed are elements of cognition among nonhuman animals? Exploring the distribution of cognitive processes in animals may provide insight into the evolution of cognition (Emery & Clayton, 2004;Gallistel, 1990). This review focuses on episodic memory (see Table 1). Students of human memory focus on episodic memory because it stores personal past experiences of an individual. In this respect, episodic memory is self-referencing. By contrast, other memory systems store facts without retaining other features that accompany memory storage. Moreover, students of human memory have been concerned with subjective experiences that are thought to accompany episodic memory retrieval in people (Tulving, 1985(Tulving, , 1987. However, documenting behavioral expression of a putative subjective experience is problematic in animals. This review advocates that it is profitable to focus on the content of episodic memories, rather than the subjective experiences that may accompany episodic memory. The central hypothesis of an animal model of episodic memory is that, at the moment of a memory assessment, the animal remembers back in time and retrieves a memory of the earlier event or episode (Crystal, 2013b(Crystal, , 2016a(Crystal, , 2016b). An important alternative explanation exists whenever the animal can solve the memory test without remembering back to the specific earlier event. In this review, I focus on judgments of familiarity as a primary nonepisodic memory alternative. According to this view, the presentation of a stimulus gives rise to a memory trace that passively decays as a function of time. Because the age of memories can be detected from a comparison of memory trace strengths, an animal may solve a new-old memory test by following a relatively simple rule such as, Choose the item that currently generates the lowest level of familiarity. Critically, an animal that uses judgments of relative familiarity need not retrieve an episodic memory of the earlier event (see details that follow). Oth...