The Generation of Time in the Hippocampal Memory System

We propose that ramping time cells in the lateral entorhinal cortex can be produced by synaptic adaptation and demonstrate this in an integrate-and-fire attractor network model. We propose that competitive networks in the hippocampal system can convert these entorhinal ramping cells into hippocampal time cells and demonstrate this in a competitive network. We propose that this conversion is necessary to provide orthogonal hippocampal time representations to encode the temporal sequence of events in hippocampal episodic memory, and we support that with analytic arguments. We demonstrate that this processing can produce hippocampal neuronal ensembles that not only show replay of the sequence later on, but can also do this in reverse order in reverse replay. This research addresses a major issue in neuroscience: the mechanisms by which time is encoded in the brain and how the time representations are then useful in the hippocampal memory of events and their order.