Is implicit memory associated with the hippocampus?

Abstract

According to the traditional memory-systems view, the hippocampus is critical during explicit (conscious) long-term memory, whereas other brain regions support implicit (nonconscious) memory. In the last two decades, some fMRI studies have reported hippocampal activity during implicit memory tasks. The aim of the present discussion paper was to identify whether any implicit memory fMRI studies have provided convincing evidence that the hippocampus is associated with nonconscious processes without being confounded by conscious processes. Experimental protocol and analysis parameters included the stimulus type(s), task(s), measures of subjective awareness, explicit memory accuracy, the relevant fMRI contrast(s) or analysis, and confound(s). A systematic review was conducted to identify implicit memory studies that reported fMRI activity in the hippocampus. After applying exclusion criteria, 13 articles remained for analysis. We found that there were no implicit memory fMRI studies where subjective awareness was absent, explicit memory performance was at chance, and there were no confounds that could have driven the observed hippocampal activity. The confounds included explicit memory (including false memory), imbalanced attentional states between conditions (yielding activation of the default-mode network), imbalanced stimuli between conditions, and differential novelty. As such, not a single fMRI study provided convincing evidence that implicit memory was associated with the hippocampus. Neuropsychological evidence was also considered, and implicit memory deficits were caused by factors known to disrupt brain regions beyond the hippocampus, such that the behavioral effects could not be attributed to this region. The present results indicate that implicit memory is not associated with the hippocampus.