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The hippocampus contains engram cells that have a privileged role in the encoding and retrieval of memories4,12. Engram cells are typically identified by expression of the activity-inducible transcription factor Fos13 during the encoding of a memory7, and their synchronous activation can trigger robust memory recall8,14. There is growing evidence that neurons defined by Fos expression carry information about context that can be used to form lasting associations, for example, between a specific environment and an aversive outcome. In addition, the hippocampus contains place cells that encode spatial maps of an environment and are thought to support spatial memories and navigation2,3,15,16.

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