Maintenance of multiple working memory items by temporal segmentation.

Experimental work based on single cell recordings supports the hypothesis that working memory representations are retained by sustained neuronal firing. While this hypothesis can account for the maintenance of a single memory item, it remains unclear how multiple working memory items are represented. This account will discuss the possible physiological mechanism responsible for the maintenance of multiple working memory items including mechanisms based on sustained firing and synaptic encoding. The focus will be on temporal segmentation by phase encoding, namely the idea that several working memory items are activated sequentially at different points in time. It has been proposed that a mechanism of nested gamma (30-80 Hz) and theta (4-8 Hz) oscillations is responsible for controlling the reactivation of the memory list. This mechanism has been shown to be compatible with multiple behavioral findings on working memory such as the data from the Sternberg experiment. The theta/gamma mechanism has also received support from a large set of electrophysiological findings, however, more experimental work is required to further substantiate or falsify the model.