Physiological patterns in the hippocampo-entorhinal cortex system.

The anatomical connectivity and intrinsic properties of entorhinal cortical neurons give rise to ordered patterns of ensemble activity. How entorhinal ensembles form, interact, and accomplish emergent processes such as memory formation is not well-understood. We lack sufficient understanding of how neuronal ensembles in general can function transiently and distinctively from other neuronal ensembles. Ensemble interactions are bound, foremost, by anatomical connectivity and temporal constraints on neuronal discharge. We present an overview of the structure of neuronal interactions within the entorhinal cortex and the rest of the hippocampal formation. We wish to highlight two principle features of entorhinal-hippocampal interactions. First, large numbers of entorhinal neurons are organized into at least two distinct high-frequency population patterns: gamma (40-100 Hz) frequency volleys and ripple (140-200 Hz) frequency volleys. These patterns occur coincident with other well-defined electrophysiological patterns. Gamma frequency volleys are modulated by the theta cycle. Ripple frequency volleys occur on each sharp wave event. Second, these patterns occur dominantly in specific layers of the entorhinal cortex. Theta/gamma frequency volleys are the principle pattern observed in layers I-III, in the neurons that receive cortical inputs and project to the hippocampus. Ripple frequency volleys are the principle population pattern observed in layers V-VI, in the neurons that receive hippocampal output and project primarily to the neocortex. Further, we will highlight how these ensemble patterns organize interactions within distributed forebrain structures and support memory formation.