Complementary learning systems.

This paper reviews the fate of the central ideas behind the complementary learning systems (CLS) framework as originally articulated in McClelland, McNaughton, and O'Reilly (1995). This framework explains why the brain requires two differentially specialized learning and memory systems, and it nicely specifies their central properties (i.e., the hippocampus as a sparse, pattern-separated system for rapidly learning episodic memories, and the neocortex as a distributed, overlapping system for gradually integrating across episodes to extract latent semantic structure). We review the application of the CLS framework to a range of important topics, including the following: the basic neural processes of hippocampal memory encoding and recall, conjunctive encoding, human recognition memory, consolidation of initial hippocampal learning in cortex, dynamic modulation of encoding versus recall, and the synergistic interactions between hippocampus and neocortex. Overall, the CLS framework remains a vital theoretical force in the field, with the empirical data over the past 15 years generally confirming its key principles.