a b s t r a c tMemory retrieval is an active process that can alter the content and accessibility of stored memories. Of potential relevance for educational practice are findings that memory retrieval fosters better retention than mere studying. This so-called testing effect has been demonstrated for different materials and populations, but there is limited consensus on the neurocognitive mechanisms involved. In this review, we relate cognitive accounts of the testing effect to findings from recent brain-imaging studies to identify neurocognitive factors that could explain the testing effect. Results indicate that testing facilitates later performance through several processes, including effects on semantic memory representations, the selective strengthening of relevant associations and inhibition of irrelevant associations, as well as potentiation of subsequent learning.

Encoding and retrieval processes enhance long-term memory performance. The efficiency of encoding processes has recently been linked to representational consistency: the reactivation of a representation that gets more specific each time an item is further studied. Here we examined the complementary hypothesis of whether the efficiency of retrieval processes also is linked to representational consistency. Alternatively, recurrent retrieval might foster representational variability-the altering or adding of underlying memory representations. Human participants studied 60 Swahili-Swedish word pairs before being scanned with fMRI the same day and 1 week later. On Day 1, participants were tested three times on each word pair, and on Day 7 each pair was tested once. A BOLD signal change in right superior parietal cortex was associated with subsequent memory on Day 1 and with successful long-term retention on Day 7. A representational similarity analysis in this parietal region revealed that beneficial recurrent retrieval was associated with representational variability, such that the pattern similarity on Day 1 was lower for retrieved words subsequently remembered compared with those subsequently forgotten. This was mirrored by a monotonically decreased BOLD signal change in dorsolateral prefrontal cortex on Day 1 as a function of repeated successful retrieval for words subsequently remembered, but not for words subsequently forgotten. This reduction in prefrontal response could reflect reduced demands on cognitive control. Collectively, the results offer novel insights into why memory retention benefits from repeated retrieval, and they suggest fundamental differences between repeated study and repeated testing.

One important distinction in psychology is between inferences based on associative memory and inferences based on analysis and rules. Much previous empirical work conceive of associative and analytical processes as two exclusive ways of addressing a judgment task, where only one process is selected and engaged at a time, in an either-or fashion. However, related work indicate that the processes are better understood as being in interplay and simultaneously engaged. Based on computational modeling and brain imaging of spontaneously adopted judgment strategies together with analyses of brain activity elicited in tasks where participants were explicitly instructed to perform similarity-based associative judgments or rule-based judgments (n = 74), we identified brain regions related to the two types of processes. We observed considerable overlap in activity patterns. The precuneus was activated for both types of judgments, and its activity predicted how well a similarity-based model fit the judgments. Activity in the superior frontal gyrus predicted the fit of a rule-based judgment model. The results suggest the precuneus as a key node for similarity-based judgments, engaged both when overt responses are guided by similarity-based and rule-based processes. These results are interpreted such that similarity-based processes are engaged in parallel to rule-based-processes, a finding with direct implications for cognitive theories of judgment.

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