The dentate gyrus as a filter or gate: a look back and a look ahead

The idea of the dentate gyrus as a gate or filter at the entrance to the hippocampus, blocking or filtering incoming excitation from the entorhinal cortex, has been an intriguing one. Here we review the historical development of the idea, and discuss whether it may be possible to be more specific in defining this gate. We propose that dentate function can be understood within a context of Hebbian association and competition: hilar mossy cells help the dentate granule cells to recognize incoming entorhinal patterns of activity (Hebbian association), after which patterns that are consistently and repetitively presented to the dentate gyrus are passed through, while random, more transient patterns are blocked (non-associative Hebbian competition). Translamellar inhibition as well as translamellar potentiation can be understood in this context. The dentate-hilar complex thus plays the role of a "pattern excluder", not a pattern completer. The unique role of pattern exclusion may explain the peculiar qualities of dentate granule cells and hilar mossy cells.