Does working memory activate the hippocampus during the late delay period?

The aim of the present discussion paper was to identify whether any fMRI studies have provided convincing evidence that the hippocampus is associated with working memory. The key outcome variable was the phase in which hippocampal activity was observed: study, early delay, late delay, and/or test. During working memory tasks, long-term memory processes can operate during the study phase, early delay phase (due to extended encoding), or test phase. Thus, working memory processes can be isolated from long-term memory processes during only the late delay period. Twenty-six working memory studies that reported hippocampal activity were systematically analyzed. Many experimental protocol and analysis parameters were considered including number of participants, stimulus type(s), number of items during the study phase, delay duration, task during the test phase, behavioral accuracy, relevant fMRI contrast(s), whether the information was novel or familiar, number of phases modeled, and whether activation timecourses were extracted. For studies that were able to identify activity in different phases, familiar information sometimes produced activity during the study phase and/or test phase, but never produced activity during the delay period. When early-delay phase and late-delay phase activity could be distinguished via modeling these phases separately or inspecting activation timecourses, novel information could additionally produce activity during the early delay phase. There was no convincing evidence of hippocampal activity during the late delay period. These results indicate that working memory does not activate the hippocampus and suggest a model of working memory where maintenance of novel information can foster long-term memory encoding.