Emma K. Bridger

The FN400 is functionally distinct from the N400

The FN400 refers to the early midfrontally-distributed difference between ERPs elicited by old and new items, which operates in a way consistent with a neural marker of familiarity-based recognition. Double dissociations between the FN400 and a later ERP index of recollection provide some of the most compelling evidence in support of dual-process models to date. It has recently been claimed, however, that there is no evidence that the FN400 is functionally distinct from the N400 index of implicit semantic priming (Voss, J., and Federmeier, K., FN400 potentials are functionally identical to N400 potentials and reflect semantic processing during recognition testing, Psychophysiology, 48, 532-546, 2011), challenging inferences made on the basis of this effect. We argue that the design employed to make this claim is flawed because it comprised a semantic priming manipulation embedded within a continuous recognition test which enabled recognition contrasts to be confounded by semantic processes in a number of ways. Here, ERPs were recorded from a design which avoided these confounds by employing a semantic priming paradigm which also served as the encoding phase for a surprise subsequent recognition test phase. An N400 effect elicited in the semantic priming task demonstrated the established centro-parietal maximum, whereas the difference between correctly responded to old and new ERPs in the recognition test was maximal over frontal sites in the same time window. When direct comparisons of the electrophysiological correlates of semantic priming and episodic recognition are recorded in a paradigm in which the two are not confounded, the FN400 reflects a qualitatively distinct effect from the N400.

Neural correlates of individual differences in strategic retrieval processing

Processes engaged when information is encoded into memory are an important determinant of whether that information will be recovered subsequently. Also influential, however, are processes engaged at the time of retrieval, and these were investigated here by using event-related potentials (ERPs) to measure a specific class of retrieval operations. These operations were revealed by contrasts between ERPs elicited by new (unstudied) test items in distinct tasks, the assumption being that these contrasts index operations that are engaged in service of retrieval and that vary according to the demands of different retrieval tasks. Specific functional accounts of this class of retrieval processing operations assume that they influence the accuracy of memory judgments, and this experiment was designed to test for the first time whether this is in fact the case. Toward this end, participants completed 2 retrieval tasks while ERPs were acquired, and the extent to which processes were engaged differentially across tasks in service of retrieval was operationalized as the magnitude of the differences between the new-item ERPs that were elicited. This measure correlated positively with response accuracy on the tasks, which provides strong evidence that this class of retrieval processing operations benefits the accuracy of memory judgments.

Semantic relations differentially impact associative recognition memory: electrophysiological evidence.

Though associative recognition memory is thought to rely primarily on recollection, recent research indicates that familiarity might also make a substantial contribution when to-be-learned items are integrated into a coherent structure by means of an existing semantic relation. It remains unclear how different types of semantic relations, such as categorical (e.g., dancer-singer) and thematic (e.g., dancer-stage) relations might affect associative recognition, however. Using event-related potentials (ERPs), we addressed this question by manipulating the type of semantic link between paired words in an associative recognition memory experiment. An early midfrontal old/new effect, typically linked to familiarity, was observed across the relation types. In contrast, a robust left parietal old/new effect was found in the categorical condition only, suggesting a clear contribution of recollection to associative recognition for this kind of pairs. One interpretation of this pattern is that familiarity was sufficiently diagnostic for associative recognition of thematic relations, which could result from the integrative nature of the thematic relatedness compared to the similarity-based nature of categorical pairs. The present study suggests that the extent to which recollection and familiarity are involved in associative recognition is at least in part determined by the properties of semantic relations between the paired associates.

More ways than one: ERPs reveal multiple familiarity signals in the word frequency mirror effect

Recent dual-process models of the word frequency mirror effect place absolute familiarity, an item׳s baseline familiarity at a given time point, as responsible for false alarm differences and recollection for hit rate differences between high and low frequency items. One of the earliest dual-process propositions, however, posits an additional relative familiarity mechanism which is sensitive to recent presentation but relative to the absolute familiarity of a particular item (Mandler, 1980). In this study, it was possible to map these three mechanisms onto known event-related potential (ERP) effects in an old/new recognition task with high and low frequency words. Contrasts between ERPs elicited by high and low frequency new items were assumed to index absolute familiarity, and the distribution of this effect from 300 to 600ms was topographically distinct from a temporally-overlapping midfrontally-distributed old/new effect which was larger for low than high frequency words, as would be expected from a relative familiarity mechanism. A later left parietal old/new effect, strongly linked to recollection, was only present for low frequency items. These frequency-sensitive amplitude differences for both old/new effects disappeared in a second recognition task in which old/new decisions were made under a time constraint, although the posterior absolute familiarity effect remained unaffected by the speeding of responses. The data support the assertion that three distinct recognition processes are affected by word frequency in recognition memory tasks, and the qualitatively distinct distributions associated with the two familiarity contrasts support the presence of two cognitively distinct familiarity mechanisms.

Requirements at retrieval modulate subsequent memory effects: An event-related potential study

Neural indices of memory formation can be acquired by contrasting activity during study for items that are remembered or forgotten on a subsequent memory test. These “subsequent memory” effects vary with the stimulus types that are encoded, how they are encoded, the correspondences between study and test materials, and the time intervals between study and test phases. We investigated whether event-related potential (ERP) subsequent memory effects also vary with the content people must retrieve. Participants saw words on the left/right side of fixation, and made a drawing difficulty or pleasantness judgment to each. In separate test phases, participants were asked to remember study screen location, or which task judgment had been made. The ERP subsequent memory effects from these two tasks were functionally distinct, demonstrating for the first time that ERP subsequent memory effects dissociate according to what people are trying to retrieve.

Nap sleep preserves associative but not item memory performance

Many studies have shown that sleep improves memory performance, and that even short naps during the day are beneficial. Certain physiological components of sleep such as spindles and slow-wave-sleep are thought to be particularly important for memory consolidation. The aim of this experiment was to reveal the role of naps for hippocampus-dependent associative memory (AM) and hippocampus-independent item memory (IM) alongside their corresponding ERP old/new effects. Participants learnt single words and word-pairs before performing an IM- and an AM-test (baseline). One group was subsequently allowed to nap (∼90min) while the other watched DVDs (control group). Afterwards, both groups performed a final IM- and AM-test for the learned stimuli (posttest). IM performance decreased for both groups, while AM performance decreased for the control group but remained constant for the nap group, consistent with predictions concerning the selective impact of napping on hippocampus-dependent recognition. Putative ERP correlates of familiarity and recollection were observed in the IM posttest, whereas only the later recollection-related effect was present in the AM test. Notably, none of these effects varied with group. Positive correlations were observed between spindle density during slow-wave-sleep and AM posttest performance as well as between spindle density during non-REM sleep and AM baseline performance, showing that successful learning and retrieval both before and after sleep relates to spindle density during nap sleep. Together, these results speak for a selective beneficial impact of naps on hippocampus-dependent memories.

Sleep spindles during a nap correlate with post sleep memory performance for highly rewarded word-pairs.

HighlightsNap sleep benefits high but not low rewarded information.Spindle density correlates with high reward memory retention post‐sleep.Spindle density correlates with overall memory performance pre‐ and post‐sleep. Abstract The consolidation of new associations is thought to depend in part on physiological processes engaged during non‐REM (NREM) sleep, such as slow oscillations and sleep spindles. Moreover, NREM sleep is thought to selectively benefit associations that are adaptive for the future. In line with this, the current study investigated whether different reward cues at encoding are associated with changes in sleep physiology and memory retention. Participants’ associative memory was tested after learning a list of arbitrarily paired words both before and after taking a 90‐min nap. During learning, word‐pairs were preceded by a cue indicating either a high or a low reward for correct memory performance at test. The motivation manipulation successfully impacted retention such that memory declined to a greater extent from pre‐ to post sleep for low rewarded than for high rewarded word‐pairs. In line with previous studies, positive correlations between spindle density during NREM sleep and general memory performance pre‐ and post‐sleep were found. In addition to this, however, a selective positive relationship between memory performance for highly rewarded word‐pairs at posttest and spindle density during NREM sleep was also observed. These results support the view that motivationally salient memories are preferentially consolidated and that sleep spindles may be an important underlying mechanism for selective consolidation.

The beneficial effect of testing: an event-related potential study

The enhanced memory performance for items that are tested as compared to being restudied (the testing effect) is a frequently reported memory phenomenon. According to the episodic context account of the testing effect, this beneficial effect of testing is related to a process which reinstates the previously learnt episodic information. Few studies have explored the neural correlates of this effect at the time point when testing takes place, however. In this study, we utilized the ERP correlates of successful memory encoding to address this issue, hypothesizing that if the benefit of testing is due to retrieval-related processes at test then subsequent memory effects (SMEs) should resemble the ERP correlates of retrieval-based processing in their temporal and spatial characteristics. Participants were asked to learn Swahili-German word pairs before items were presented in either a testing or a restudy condition. Memory performance was assessed immediately and 1-day later with a cued recall task. Successfully recalling items at test increased the likelihood that items were remembered over time compared to items which were only restudied. An ERP subsequent memory contrast (later remembered vs. later forgotten tested items), which reflects the engagement of processes that ensure items are recallable the next day were topographically comparable with the ERP correlate of immediate recollection (immediately remembered vs. immediately forgotten tested items). This result shows that the processes which allow items to be more memorable over time share qualitatively similar neural correlates with the processes that relate to successful retrieval at test. This finding supports the notion that testing is more beneficial than restudying on memory performance over time because of its engagement of retrieval processes, such as the re-encoding of actively retrieved memory representations.

Is faster better? Effects of response deadline on ERP correlates of recognition memory in younger and older adults

Aging studies generally suggest that recollection is impaired whereas familiarity-based recognition remains relatively preserved in healthy older adults. The present event-related potential (ERP) study explores whether age-related impairments in recognition memory can be reduced under conditions in which recognition decisions are primarily driven by familiarity. Old and young adults performed an item recognition task with perceptually rich visual stimuli. A response deadline procedure was employed following previous studies which have shown that limiting response times attenuates recollection but leaves familiarity relatively unaffected. Age effects on memory performance were large in the non-speeded response condition in which recollection contributes to performance. When response time was limited, performance differences between groups were negligible. In the non-speeded condition the ERP correlate of recollection was not detectable in old adults. Conversely, in the speeded condition ERP correlates of familiarity were obtained in both age groups, though attenuated for old adults. For old adults in the speeded condition a temporally extended posterior negativity was obtained which was more pronounced for low performing participants. The results suggest that even though the neural generators of the familiarity signal degrade with age, familiarity is an important contributor to recognition memory in older adults and can lead to a disproportional benefit in memory in conditions designed to specifically enhance familiarity-based responding.

Age effects on associative memory for novel picture pairings

Normal aging is usually accompanied by greater memory decline for associations than for single items. Though associative memory is generally supported by recollection, it has been suggested that familiarity can also contribute to associative memory when stimuli can be unitized and encoded as a single entity. Given that familiarity remains intact during healthy aging, this may be one route to reducing age-related associative deficits. The current study investigated age-related differences in associative memory under conditions that were expected to differentially promote unitization, in this case by manipulating the spatial arrangement of two semantically unrelated objects positioned relative to each other in either spatially implausible or plausible orientations. Event-related potential (ERP) correlates of item and associative memory were recorded whilst younger and older adults were required to discriminate between old, recombined and new pairs of objects. These ERP correlates of item and associative memory did not vary with plausibility, whereas behavioral measures revealed that both associative and item memory were greater for spatially plausible than implausible pair arrangements. Contrary to predictions, older adults were less able to take advantage of this memory benefit than younger participants. Potential reasons for this are considered, and these are informed by those lines of evidence which indicate older participants were less sensitive to the bottom-up spatial manipulation employed here. It is recommended that future strategies for redressing age-related associative deficits should take account of the aging brains increasing reliance on pre-existing semantic associations.