Event-related potentials during conscious and automatic memory retrieval

The effects of study-test lags of between 0 and 32 items on conscious (C) and automatic (A) memory processes in a running word-completion task were investigated with event-related potentials (ERPs). The process dissociation procedure (PDP) can distinguish between C and A contributions to memory by comparing performance when subjects respond with either an old item (inclusion) or a new item (exclusion). C can be estimated by subtracting the probability of an intrusion of an old item during the exclusion task (due to A without C) from the probability of correctly producing an old item during the inclusion task (due to C and/or A). The behavioral results showed that C was stronger when the test item followed the studied word in the next trial or after a lag of one stimulus. The strength of A did not vary with lag. The ERP waveforms contained a broad parietal positive wave between 300 and 800 ms. This parietal wave distinguished between correctly recalled old and new words. The early portion of this old-new effect was significantly affected by lag. Subtracting waveforms to obtain a measure of C revealed an effect in the later portion of this wave, lateralized over the left hemisphere. A sustained frontal negativity occurred during all recordings and was larger during conscious retrieval. There was no consistent ERP effect related to automatic memory retrieval.     

Event-related potential correlates of the retrieval of emotional and nonemotional context

In two experiments, we examined event-related potentials (ERPs) elicited in an old/new recognition memory test by emotionally neutral visual objects that, at encoding, had been associated with neutrally, negatively, or positively valenced background contexts. In Experiment 2, subjects also judged the context in which the item had been studied. In Experiment 1, "left parietal" old/new ERP effects were elicited by correctly recognized items. Items encoded in emotional contexts, but not those studied in neutral contexts, elicited additional effects early in the recording epoch over lateral temporal scalp and, later, over left temporo-frontal scalp. In Experiment 2, "left parietal" and "right frontal" ERP effects were elicited by recognized items that attracted correct source judgments. Additional effects, an early lateral temporal positivity and a late-onset, left-sided positivity, were elicited by items studied in emotionally valenced contexts and attracting correct source judgments. Together, the findings indicate that retrieval processing is influenced by the emotional valence of the context in which an item is encoded, regardless of whether contextual information is task relevant.     

Recognition memory for emotionally negative and neutral words: an ERP study

Scalp recorded event-related potentials were used to investigate the neural activity elicited by emotionally negative and emotionally neutral words during the performance of a recognition memory task. Behaviourally, the principal difference between the two word classes was that the false alarm rate for negative items was approximately double that for the neutral words. Correct recognition of neutral words was associated with three topographically distinct ERP memory 'old/new' effects: an early, bilateral, frontal effect which is hypothesised to reflect familiarity-driven recognition memory; a subsequent left parietally distributed effect thought to reflect recollection of the prior study episode; and a late onsetting, right-frontally distributed effect held to be a reflection of post-retrieval monitoring. The old/new effects elicited by negative words were qualitatively indistinguishable from those elicited by neutral items and, in the case of the early frontal effect, of equivalent magnitude also. However, the left parietal effect for negative words was smaller in magnitude and shorter in duration than that elicited by neutral words, whereas the right frontal effect was not evident in the ERPs to negative items. These differences between neutral and negative words in the magnitude of the left parietal and right frontal effects were largely attributable to the increased positivity of the ERPs elicited by new negative items relative to the new neutral items. Together, the behavioural and ERP findings add weight to the view that emotionally valenced words influence recognition memory primarily by virtue of their high levels of 'semantic cohesion', which leads to a tendency for 'false recollection' of unstudied items.     

The effect of item sequence on brain activity during recognition memory

Two basic types of experimental designs are available to measure the brain indices of successful recognition memory. Both designs have in common that brain activity elicited by correctly recognized studied (old) items is compared to brain activity elicited by correctly rejected new items. In blocked designs, the two item types are presented in separate lists. In random-mixed designs, on the other hand, they are randomly intermixed within the same list. Early studies reported that in random-mixed designs, brain activity can be affected by the sequence in which different types of items are presented. Here, we investigated to what extent such sequence effects also occur in random-mixed designs of recognition memory. Event-related potentials (ERPs) were obtained while subjects made recognition judgments on pseudorandom sequences of visually presented new and old words. Changes in the sequence from one item-type to another (change from old to new and from new to old) were associated with a prominent P300 potential for new words following old words but not vice versa. Subjects with larger P300 potentials made less false alarms to such new words. Successive presentations of the same item-type (sequence 'blocks': three old words in a row and three new words in row) were associated with a left parietal positive slow shift for successive old words but not for successive new words. The data suggest that new words require extensive context-updating if they are presented within the context generated by preceding old words. Due to these sequence effects, the present data question the assumption that random-mixed designs are generally favorable over block designs.     

Strategic retrieval in a reality monitoring task

Strategic recollection refers to control processes that allow the retrieval of information that is relevant for a specific situation. These processes can be studied in memory exclusion tasks, which require the retrieval of particular kinds of episodic information. In the current study, we investigated strategic recollection in reality monitoring by event-related potentials (ERPs). Participants studied object words, followed by a picture of the denoted object (perceive condition) or followed by the instruction to imagine such a picture (imagine condition). At test, subjects had to identify words of one study condition and to reject words of the second study condition together with newly presented items. Data analysis showed that object names were better identified when items of the perceive condition were targeted. In this test condition, a left parietal old/new effect (the ERP correlate of recollection) was observed only in response to targets. In contrast, both targets and nontargets elicited this old/new effect when items of the imagine condition were targeted. The magnitude of the left parietal old/new effect to nontargets in this condition (but no other left parietal old/new effect) correlated positively with the discrimination indices of both test conditions. In addition, ERPs to targets and nontargets differed at right frontal electrode sites at longer latencies (1500-1800 ms), with more positive ERPs for targets. Findings indicate that subjects retrieved nontarget information in the more difficult task condition, while they relied on target information alone in the less difficult task. This kind of strategic retrieval was not mirrored in other old/new effects. The correlation between the left parietal old/new effect for nontargets in the imagined item target condition and the discrimination indices of both conditions may indicate that the ease of nontarget retrieval, rather than the difficulty of target retrieval, increases the likelihood that nontarget information is actually retrieved.     

Electrophysiological correlates of emotion-induced recognition bias

The question of how emotions influence recognition memory is of interest not only within basic cognitive neuroscience but from clinical and forensic perspectives as well. Emotional stimuli can induce a "recognition bias" such that individuals are more likely to respond "old" to a negative item than to an emotionally neutral item, whether the item is actually old or new. We investigated this bias using event-related brain potential (ERP) measures by comparing the processing of words given "old" responses with accurate recognition of old/new differences. For correctly recognized items, the ERP difference between old items (hits) and new items (correct rejections, CR) was largely unaffected by emotional valence. That is, regardless of emotional valence, the ERP associated with hits was characterized by a widespread positivity between 300 and 700 msec relative to that for CRs. By contrast, the analysis of ERPs to old and new items that were judged "old" (hits and false alarms [FAs], respectively) revealed a differential effect of valence by 300 msec: Neutral items showed a large old/new difference over prefrontal sites, whereas negative items did not. These results are the first clear demonstration of response bias effects on ERPs linked to recognition memory. They are consistent with the idea that frontal cortex areas may be responsible for relaxing the retrieval criterion for negative stimuli so as to ensure that emotional events are not as easily "missed" or forgotten as neutral events.     

Event-related potential studies of associative recognition and recall: electrophysiological evidence for context dependent retrieval processes

To investigate the neural correlates of episodic recollection the ERP correlates of memory for new associations (recently studied novel word pairs) were investigated using two tasks, associative recognition and associative recall. For the recognition task subjects discriminated old from new word pairs and, for pairs judged old, reported whether the pairs were intact or recombined (compared to at study). For the recall task, subjects discriminated old from new words and, for each word judged old, reported its study associate. ERPs were recorded at test from 25 scalp electrodes, with a 1944-ms recording epoch. In Experiment 1, the tasks were randomly interleaved. Consistent with previous findings, relative to the ERPs for correctly classified new items, the ERP correlates of successful associative recognition consisted of a sustained left parietal positivity, and two frontal positivities, one early and bilateral, the other occurring later and showing a right-sided maximum. In contrast to previous findings, successful associative recall elicited similar effects to those found for recognition. Topographic analyses revealed that the distribution of these retrieval-related ERP effects were similar across the two tasks, suggesting that the recognition and recall of associative information gives rise to activity in overlapping, if not the same, neural populations. In Experiment 2 the tasks were blocked. In contrast to the findings of Experiment 1, successful associative recall elicited left parietal and late onsetting right frontal positivities, in the absence of the early bilateral frontal positivity. This finding suggests that frontally-distributed memory-related ERP effects are both neurally and functionally dissociable. Specifically, we argue that the functional significance of the early frontally distributed ERP effect cannot be accounted for by the 'post-retrieval processing' hypothesis that is taken to account for the late right frontal effect, suggesting that episodic recollection itself is neither neurally nor functionally homogenous.     

Effects of attention and confidence on the hypothesized ERP correlates of recollection and familiarity

Dual-process theories suggest that recognition memory is determined by two separate processes: familiarity and recollection. Experiment 1 behaviorally replicated past studies using the remember/know procedure to indicate that the amount of attention devoted to study influences both recollection and familiarity, but recollection more strongly. Experiments 1 and 2 assessed the effects of attention on two ERP components that have been hypothesized to be related to familiarity (FN400 old/new effect, 300-500 ms, anterior) and recollection (parietal old/new effect, 400-800 ms, posterior). Parietal old/new effects were reduced by divided attention, but FN400 old/new effects were not. Parietal ERPs (400-800 ms) in experiment 2 increased with confidence in recognizing old items, but not new items. These results support the hypothesis that the parietal old/new effect is related to recollection.     

Predictive validity of event-related potentials (ERPs) in relation to the directed forgetting effects.

OBJECTIVE: A directed forgetting paradigm (word method) was used to assess the relationships between the event-related potentials (ERPs) recorded during the study phase and the subsequent forgetting effects. METHODS: In the study phase 100 words were presented each followed by either an instruction to remember (R) or to forget (F). Then these 100 words, together with another 100 new words, were presented and subjects had to perform an old/new decision task. The forgetting effect was considered as the difference between the rate of R- and F-words correctly recognised. Based on this difference, the whole sample was divided into two 10-subject groups with high and low forgetting effect. RESULTS: From 300 to 600 msec post-instruction differences in ERPs between R and F instructions were similar in both groups. In contrast, in the 100-300 msec epoch, ERP differences only appeared in the high-effect group and consisted of an enhanced positive activity elicited by F instruction at frontal and prefrontal areas and a larger positivity associated to R instruction at parietal area. CONCLUSIONS: The findings suggest that the processing of the word is kept on stand-by until the instruction is provided. Instruction to remember would reactivate this processing, which would mainly involve the parietal area, whereas instruction to forget appears to trigger frontal and prefrontal mechanisms trying to inhibit such processing.     

Event-related brain potentials during auditory and visual word recognition memory tasks

Event-related brain potentials (ERPs) recorded during presentation of a series of words or pictures show enhanced positivity between 300 and 800 ms after presentation of repeated items. However, little attention has been directed to the characterization of this ERP recognition memory effect using auditory stimuli. The present study directly compared the ERP 'old/new effect' for words presented in the visual and auditory modalities. Nose-referenced ERPs were recorded from 30 electrode sites while participants (N=16) were engaged in visual and auditory continuous word recognition memory tasks. Spatially and temporally overlapping ERP components were identified and measured by covariance-based principal components analysis. The expected old/new effect was observed in both modalities, with a comparable time course peaking at 560 ms, but having a more anterior scalp topography for visual items. This suggests a common cognitive process (i.e. successful retrieval of information from memory) associated with separable neural generators in each modality. Despite this temporal synchronization, the old/new effect overlapped ERP components having distinct scalp topographies (N2) or peak latencies (P3) for each modality. The positive-going old/new effect was preceded by an earlier negativity peaking at 370 ms that was greater across modalities for old than new words, likely reflecting semantic processing aspects of word recognition memory. A late (beyond 900 ms), broadly-distributed negativity was also greater for old than new words, prolonged for auditory items, and may represent activity of a post-retrieval process.     

Using ERPs to dissociate recollection from familiarity in picture recognition

Dual process theories posit that separate recollection and familiarity processes contribute to recognition memory. Previous research, testing recognition memory for words, indicates that event-related brain potentials (ERPs) can be used to dissociate recollection from familiarity. It has been hypothesized that the FN400 ERP old/new effect (300-500 ms) varies with stimulus familiarity, but the parietal ERP old/new effect (400-800 ms) varies with recollection. The results reported here are consistent with this hypothesis, extending it to the recognition of pictures when subjects had to discriminate between studied pictures, highly familiar lures (mirror-reversals of studied pictures), and new pictures. Furthermore, the parietal old/new effect showed significant recollection-related differences only for subjects with good behavioral discrimination between studied items and similar lures.     

Functional significance of retrieval-related activity in lateral parietal cortex: Evidence from fMRI and ERPs

The present study addressed the question whether neural activity in left lateral parietal cortex is modulated by amount of information recollected. In two experiments (one using fMRI and the other ERPs), subjects first studied pairs of pictures presented for either 1 or 6 s. They then performed a standard "Remember/Know" recognition memory test in which the old items comprised one of the pictures from each studied pair. In both experiments, a surprise posttest indicated that subjects recollected more details about the study presentation of the items presented for the longer duration. In the fMRI experiment, recollection- and familiarity-based recognition elicited activity in distinct cortical networks. Additionally, recollection-related activity in left inferior parietal cortex was of greater magnitude for test items presented for 6 s than for 1 s. In the ERP study the "left-parietal old/new effect"-a putative correlate of successful recollection-was likewise modulated by amount of information retrieved. Together, these findings provide further support for dual-process models of recognition memory and add weight to the proposal that retrieval-related activity in left inferior parietal cortex reflects processes supporting the online representation of retrieved episodic information.     

Electrophysiological evidence for the modulation of retrieval orientation by depth of study processing

Event-related potentials (ERPs) were employed to investigate whether brain activity elicited by retrieval cues in a memory test varies according to the encoding task undertaken at study. Two recognition memory test blocks were administered, preceded, in one case, by a "shallow" study task (alphabetic judgement) and, in the other case, by a "deep" task (sentence generation). ERPs elicited by the new words in each test block differed, the ERPs elicited in the block following the shallow study task exhibiting the more positive-going waveforms. This finding was taken as evidence that subjects adopt different "retrieval sets" when attempting to retrieve items that had been encoded in terms of alphabetic versus semantic attributes. Differences between the ERPs elicited by correctly classified old and new words (old/new effects) also varied with encoding task. The effects for deeply studied words resembled those found in previous ERP studies of recognition memory, whereas old/new effects for shallowly studied words were confined to a late-onsetting, right frontal positivity. Together, the findings indicate that the depth of study processing influences two kinds of memory-related neural activity, associated with memory search operations, and the processing of retrieved information, respectively.     

Emotion/cognition-coupling in word recognition memory of depressive patients: an event-related potential study

Depressive patients show deficits in memory functions. However, the underlying mechanisms remain unclear. Experiments with a special emphasis on the link between emotion and cognition appear challenging. The aim of this study is to investigate the influence of the emotional content of words on memory in non-medicated depressive patients (n=11) compared with a control group (n=11) utilizing event-related brain potentials (ERPs). In a continuous word recognition paradigm brain responses to repeated items are characterized by more positive waveforms of ERPs. This recognition effect ('old/new effect') has been shown to be sensitive to parameters relevant for memory processing. For the purpose of this ERP experiment visually presented words were classified into three different categories of emotional content. The ERPs for the correctly detected 'old' (repeated) words showed an increased positivity beginning approximately 250 ms post-stimulus, concurring with a good recognition performance. In addition, old/new effect and behavioral data were sensitive to words' different emotional connotations in the control group. In contrast, the depressive patients performed worse and showed no significant old/new effect. Nevertheless, their recognition performance was also enhanced by the emotional content. Furthermore, a differential effect of the emotional content on frontal ERPs was found between groups. In contrast to the control group, a reduced old/new effect indicates a reduced working memory capacity in the moderately depressed patients. This is suggested to be partially due to changes of the emotion/cognition coupling related to ruminations with preferably negative emotional connotation. However, the emotional content also affects recognition performance in the depressive patients.     

ERP 'old/new' effects: memory strength and decisional factor(s)

Event-related potentials (ERPs) were recorded while subjects made old/new recognition judgments on new unstudied words and old words which had been presented at study either once (‘weak’) or three times (‘strong’). The probability of an ‘old’ response was significantly higher for strong than weak words and significantly higher for weak than new words. Comparisons were made initially between ERPs to new, weak and strong words, and subsequently between ERPs associated with six strength-by-response conditions. The N400 component was found to be modulated by memory trace strength in a graded manner. Its amplitude was most negative in new word ERPs and most positive in strong word ERPs. This ‘N400 strength effect’ was largest at the left parietal electrode (in ear-referenced ERPs). The amplitude of the late positive complex (LPC) effect was sensitive to decision accuracy (and perhaps confidence). Its amplitude was larger in ERPs evoked by words attracting correct versus incorrect recognition decisions. The LPC effect had a left>right, centro-parietal scalp topography (in ear-referenced ERPs). Hence, whereas, the majority of previous ERP studies of episodic recognition have interpreted results from the perspective of dual-process models, we provide alternative interpretations of N400 and LPC old/new effects in terms of memory strength and decisional factor(s).     

ERP old/new effects at different retention intervals in recency discrimination tasks

Recognition memory studies have suggested that event-related brain potentials (ERPs) may tap into several different memory processes. In particular, two ERP components have been hypothesized as related to familiarity (FN400 old/new effect, 300-500 ms, anterior) and recollection processes (parietal old/new effect, 400-800 ms, posterior). The functional significance of the FN400 old/new effect is uncertain because similar old/new differences have been shown to disappear at moderately long retention intervals. The present study investigated the effects of retention interval (34 min, 39 min, or 1 day) on the FN400 and parietal old/new effects in two different recency discrimination tasks. The results suggest that the FN400 old/new effect can be maintained across 1-day retention intervals, so it may index brain processes capable of contributing to long-term memory.     

The development of control processes supporting source memory discrimination as revealed by event-related potentials

Improvement in source memory performance throughout childhood is thought to be mediated by the development of executive control. As postretrieval control processes may be better time-locked to the recognition response rather than the retrieval cue, the development of processes underlying source memory was investigated with both stimulus- and response-locked event-related potentials (ERPs). These were recorded in children, adolescents, and adults during a recognition memory exclusion task. Green- and red-outlined pictures were studied, but were tested in black outline. The test requirement was to endorse old items shown in one study color ("targets") and to reject new items along with old items shown in the alternative study color ("nontargets"). Source memory improved with age. All age groups retrieved target and nontarget memories as reflected by reliable parietal episodic memory (EM) effects, a stimulus-locked ERP correlate of recollection. Response-locked ERPs to targets and nontargets diverged in all groups prior to the response, although this occurred at an increasingly earlier time point with age. We suggest these findings reflect the implementation of attentional control mechanisms to enhance target memories and facilitate response selection with the greatest and least success, respectively, in adults and children. In adults only, response-locked ERPs revealed an early-onsetting parietal negativity for nontargets, but not for targets. This was suggested to reflect adults' ability to consistently inhibit prepotent target responses for nontargets. The findings support the notion that the development of source memory relies on the maturation of control processes that serve to enhance accurate selection of task-relevant memories.     

The relationship between electrophysiological correlates of recollection and amount of information retrieved

The electrophysiological correlates of recollection were investigated with a modified Remember/Know task in which subjects signaled whether they fully or partially recollected visual object information in each study episode. A positive-going ERP deflection--the left parietal old/new effect--was sensitive to the amount of information recollected, demonstrating greater amplitude when elicited by test items associated with full relative to partial recollection. These findings support prior proposals that the left parietal ERP old/new effect is sensitive to the amount of information recollected from episodic memory. An early-onsetting (ca. 150 ms), left frontal old/new effect differentiated items accorded correct old versus correct new responses regardless of whether the items were endorsed as familiar or recollected. This finding extends the range of circumstances under which early, frontally distributed old/new effects occur, and adds weight to previous suggestions that these effects are a neural correlate of familiarity-driven recognition memory.     

An electrophysiological test of directed forgetting: the role of retrieval inhibition

A central issue in the research of directed forgetting is whether the differential memory performance for to-be-remembered (TBR) and to-be-forgotten (TBF) items is solely due to differential encoding or whether retrieval inhibition of TBF items plays an additional role. In this study, recognition-related event-related brain potentials (ERPs) were used to examine this issue. The spatio-temporal distributions of the old/new ERP effects obtained in Experiment 1 that employed a directed forgetting paradigm were compared with those recorded in Experiment 2 in which the level of processing was manipulated. In Experiment 1, participants were instructed to remember or to forget words by means of a cue presented after each word. ERPs recorded in the recognition test revealed early phasic frontal and parietal old/new effects for TBR items, whereas TBF items elicited only a frontal old/new effect. Moreover, a late right-frontal positive slow wave was more pronounced for TBF items, suggesting that those items were associated with a larger amount of post-retrieval processing. In Experiment 2, the same cueing method and the same stimulus materials were used, and memory encoding was manipulated by cueing participants to process the words either deeply or shallowly. Both deeply and shallowly encoded items elicited phasic frontal and parietal old/new effects followed by a late right-frontal positive slow wave. However, in contrast to TBR and TBF items, these effects differed only quantitatively. The results suggest that differential encoding alone cannot account for the effects of directed forgetting. They are more consistent with the view that items followed by an instruction to forget become inhibited and less accessible, and, therefore, more difficult to retrieve.     

Retrieval orientation and the control of recollection

Event-related potentials (ERPs) were employed to investigate whether recognition test items are processed differently according to whether they are used to probe memory for previously studied words or pictures. In each of two study-test blocks, subjects encoded a mixed list of words and pictures, and then performed a recognition memory task with words as the test items. In one block, the requirement was to respond positively to test items corresponding to studied words, and to reject both new items and items corresponding to the studied pictures. In the other block, positive responses were made to test items corresponding to pictures, and items corresponding to words were classified along with the new items. ERPs elicited during the test phase by correctly classified new items differed according to whether words or pictures were the sought-for modality. This finding was interpreted as a neural correlate of the different retrieval orientations adopted when searching memory for words versus pictures. Relative to new items, correctly classified items studied in both target modalities elicited robust, positive-going "old/new" effects. When pictures were targets, test items corresponding to studied words also elicited large effects. By contrast, when words were targets, old/new effects were absent for the items corresponding to studied pictures. These findings were interpreted as evidence that, in some circumstances, adoption of an appropriate retrieval orientation permits retrieval cues to be employed with a high degree of specificity.