In this review, we seek to identify and connect key overarching principles - principles that seem to largely hold across neural regions and tasks - that lead to neurons participating across multiple memory stages and forming a cellular substrate of memory. In this regard, significant progress has been made in our understanding memory stages at the level of ‘engram neurons’ – that is, neurons that mediate a particular memory across stages. ![]() As such, mechanistically interpreting memory in the brain is facilitated by understanding the neural underpinnings of each of these stages independently, as well as how these neural elements interrelate across stages. Memory is often conceptualized as a multi-staged process that includes encoding, consolidation, retrieval, and forgetting. Memory can be defined as an experience-dependent alteration in behavior that persists beyond the environmental stimuli that produced it. In collection, our review synthesizes general principles of the engram across memory stages, and describes future avenues to further understand the dynamic engram. Motivated by recent experimental results across these four sections, we conclude by proposing some conceptual extensions to the traditional view of the engram, including broadening the view of cell-type participation within engrams and across memory stages. Fourth, we describe neurobiological mechanisms of forgetting, and how these mechanisms can counteract engram properties established during memory encoding, consolidation, and retrieval. Third, we describe how these changes during encoding and consolidation guide neural reactivation during retrieval, and facilitate memory recall. Second, we highlight how these encoding neurons preferentially participate in synaptic- and systems-level consolidation of memory. To do so, we first describe how cell-intrinsic properties shape the initial emergence of the engram at memory encoding. Here, we integrate recent progress in the engram field to illustrate how engram neurons transform across the “lifespan” of a memory - from initial memory encoding, to consolidation and retrieval, and ultimately to forgetting. ![]() Tremendous strides have been made in our understanding of the neurobiological substrates of memory – the so-called memory “engram”.
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