Prestimulus subsequent memory effects (preSMEs)—differences in neural activity elicited by a task cue at encoding that are predictive of later memory performance—are thought to reflect differential engagement of preparatory processes that benefit episodic memory encoding. We investigated age differences in preSMEs indexed by differences in ERP amplitude just before the onset of a study item. Young and older adults incidentally encoded words for a subsequent memory test. Each study word was preceded by a task cue that signaled a judgment to perform on the word. Words were presented for either a short (300 msec) or long (1000 msec) duration with the aim of placing differential benefits on engaging preparatory processes initiated by the task cue. ERPs associated with subsequent successful and unsuccessful recollection, operationalized here by source memory accuracy, were estimated time-locked to the onset of the task cue. In a late time window (1000–2000 msec after onset of the cue), young adults demonstrated frontally distributed preSMEs for both the short and long study durations, albeit with opposite polarities in the two conditions. This finding suggests that preSMEs in young adults are sensitive to perceived task demands. Although older adults showed no evidence of preSMEs in the same late time window, significant preSMEs were observed in an earlier time window (500–1000 msec) that was invariant with study duration. These results are broadly consistent with the proposal that older adults differ from their younger counterparts in how they engage preparatory processes during memory encoding.

Existing data from noninvasive studies have led researchers to posit that the posterior cingulate cortex (PCC) supports mnemonic processes: It exhibits degeneration in memory disorders, and fMRI investigations have demonstrated memory-related activation principally during the retrieval of memory items. Despite these data, the role of the PCC in episodic memory has received only limited treatment using the spatial and temporal precision of intracranial EEG, with previous analyses focused on item retrieval. Using data gathered from 21 human participants who underwent stereo-EEG for seizure localization, we characterized oscillatory patterns in the PCC during the encoding and retrieval of episodic memories. We identified a subsequent memory effect during item encoding characterized by increased gamma band oscillatory power and a low-frequency power desynchronization. Fourteen participants had stereotactic electrodes located simultaneously in the hippocampus and PCC, and with these unique data, we describe connectivity changes between these structures that predict successful item encoding and that precede item retrieval. Oscillatory power during retrieval matched the pattern we observed during encoding, with low-frequency (below 15 Hz) desynchronization and a gamma band (especially high gamma, 70–180 Hz) power increase. Encoding is characterized by synchrony between the hippocampus and PCC, centered at 3 Hz, consistent with other observations of properties of this oscillation akin to those for rodent theta activity. We discuss our findings in light of existing theories of episodic memory processing, including the information via desynchronization hypothesis and retrieved context theory, and examine how our data fit with existing theories for the functional role of the PCC. These include a postulated role for the PCC in modulating internally directed attention and for representing or integrating contextual information for memory items.

fMRI was employed to assess whether the contents of recollection vary according to retrieval goal. At study, visually presented words were superimposed on urban or rural scenes or a gray background. The word–background pairs were presented in one of three spatial locations. During a scanned test phase, studied and unstudied words were presented. Two different source memory tasks were randomly interleaved. In the “background” task, the requirement was to judge whether the word had been presented against one of the two classes of scene, as opposed to the alternate class or the gray background. In the “location” task, discrimination was between words presented in one of the two lateral locations and words presented in either of the alternate locations. In both tasks, unstudied words required a separate response. In the background task, words studied against scenes elicited greater activity in parahippocampal and retrosplenial cortex than did words studied against the gray background, consistent with prior reports of scene reinstatement effects. Reinstatement effects were also evident in the location task. Relative to the background task, however, the effects were attenuated in parahippocampal cortex. In other regions, including medial prefrontal and posterior cingulate cortex, activity elicited in the location task by items associated with scenes was lower than that elicited by items presented on the gray background. The findings are interpreted as evidence that contextual retrieval is partially modulated by retrieval goal.

This study used event-related fMRI to examine the impact of the adoption of different retrieval orientations on the neural correlates of recollection. In each of two study–test blocks, participants 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. On the basis of previous ERP findings, we predicted that in the word task, recollection-related effects would be found for target information only. This prediction was fulfilled. In both tasks, targets elicited the characteristic pattern of recollection-related activity. By contrast, nontargets elicited this pattern in the picture task, but not in the word task. Importantly, the left angular gyrus was among the regions demonstrating this dissociation of nontarget recollection effects according to retrieval orientation. The findings for the angular gyrus parallel prior findings for the “left-parietal” ERP old/new effect and add to the evidence that the effect reflects recollection-related neural activity originating in left ventral parietal cortex. Thus, the results converge with the previous ERP findings to suggest that the processing of retrieval cues can be constrained to prevent the retrieval of goal-irrelevant information.

ERPs were recorded from samples of young (18–29 years) and older (63–77 years) participants while they performed a modified “remember–know” recognition memory test. ERP correlates of familiarity-driven recognition were obtained by contrasting the waveforms elicited by unrecollected test items accorded “confident old” and “confident new” judgments. Correlates of recollection were identified by contrasting the ERPs elicited by items accorded “remember” and confident old judgments. Behavioral analyses revealed lower estimates of both recollection and familiarity in older participants than in young participants. The putative ERP correlate of recollection—the “left parietal old–new effect”—was evident in both age groups, although it was slightly but significantly smaller in the older sample. By contrast, the putative ERP correlate of familiarity—the “midfrontal old–new effect”—could be identified in young participants only. This age-related difference in the sensitivity of ERPs to familiarity was also evident in subgroups of young and older participants, in whom familiarity-based recognition performance was equivalent. Thus, the inability to detect a reliable midfrontal old–new effect in older participants was not a consequence of an age-related decline in the strength of familiarity. These findings raise the possibility that familiarity-based recognition memory depends upon qualitatively different memory signals in older and young adults.

Recall of a studied item and retrieval of its encoding context (source memory) both depend on recollection of qualitative information about the study episode. This study investigated whether recall and source memory engage overlapping neural regions. Participants ( n = 18) studied a series of words, which were presented either to the left or right of fixation. fMRI data were collected during a subsequent test phase in which three-letter word-stems were presented, two thirds of which could be completed by a study item. Instructions were to use each stem as a cue to recall a studied word and, when recall was successful, to indicate the word's study location. When recall failed, the stem was to be completed with the first word to come to mind. Relative to stems for which recall failed, word-stems eliciting successful recall were associated with enhanced activity in a variety of cortical regions, including bilateral parietal, posterior midline, and parahippocampal cortex. Activity in these regions was enhanced when recall was accompanied by successful rather than unsuccessful source retrieval. It is proposed that the regions form part of a “recollection network” in which activity is graded according to the amount of information retrieved about a study episode.

The neural correlates of the encoding of associations between pairs of words, pairs of pictures, and word–picture pairs were compared. The aims were to determine, first, whether the neural correlates of associative encoding vary according to study material and, second, whether encoding of across- versus within-material item pairs is associated with dissociable patterns of hippocampal and perirhinal activity, as predicted by the “domain dichotomy” hypothesis of medial temporal lobe function. While undergoing fMRI scanning, subjects ( n = 24) were presented with the three classes of study pairs, judging which of the denoted objects fit into the other. Outside of the scanner, subjects then undertook an associative recognition task, discriminating between intact study pairs, rearranged pairs comprising items that had been presented on different study trials, and unstudied item pairs. The neural correlates of successful associative encoding—subsequent associative memory effects—were operationalized as the difference in activity between study pairs correctly judged intact versus pairs incorrectly judged rearranged on the subsequent memory test. Pair type–independent subsequent memory effects were evident in the left inferior frontal gyrus (IFG) and the hippocampus. Picture–picture pairs elicited material-selective effects in regions of fusiform cortex that were also activated to a greater extent on picture trials than on word trials, whereas word–word pairs elicited material-selective subsequent memory effects in left lateral temporal cortex. Contrary to the domain-dichotomy hypothesis, neither hippocampal nor perirhinal subsequent memory effects differed depending on whether they were elicited by within- versus across-material study pairs. It is proposed that the left IFG plays a domain-general role in associative encoding, that associative encoding can also be facilitated by enhanced processing in material-selective cortical regions, and that the hippocampus and perirhinal cortex contribute equally to the formation of inter-item associations, regardless of whether the items belong to the same or to different processing domains.

fMRI (1.5 mm isotropic voxels) was employed to investigate the relationship between hippocampal activity and memory strength in a continuous recognition task. While being scanned, subjects were presented with colored photographs that each appeared on four occasions. The requirements were to make one response when an item was presented for the first or the third time and to make a different response when an item appeared for the second or the fourth time. Consistent with prior findings, items presented for the first time elicited greater hippocampal and parahippocampal activity than repeated items. The activity elicited by repeated items declined linearly as a function of number of presentations (“graded” new > old effects). No medial-temporal lobe regions could be identified where activity elicited by repeated items exceeded that for new items or where activity elicited by repeated items increased with number of presentations. These findings are inconsistent with the proposal that retrieval-related hippocampal activity is positively correlated with memory strength. We also identified graded new > old effects in several cortical regions outside the medial-temporal lobe, including the left retrosplenial/posterior cingulate cortex and the right lateral occipito-temporal cortex. By contrast, graded old > new effects were evident in bilateral mid-intraparietal sulcus and precuneus.

The electrophysiological correlates of retrieval orientation—the differential processing of retrieval cues according to the nature of the sought-for information—were investigated in healthy young (18–20 years old) and older (63–77 years old) adults. In one pair of study–test cycles, subjects studied either words or pictures presented in one of two visually distinct contexts, and then performed a yes/no recognition task with words as test items. In another pair of study–test cycles, subjects again made recognition judgments, but were required, in addition, to signal the study context for each item judged “old.” Young subjects' event-related potentials (ERPs) for new (unstudied) test items were more negative-going when the study material was pictures rather than words, and this effect varied little between the two retrieval tasks. Replicating a previous report [Morcom, A. M., & Rugg, M. D. Effects of age on retrieval cue processing as revealed by ERPs. Neuropsychologia, 42, 1525–1542, 2004], the effects of study material on the ERPs of the older subjects were attenuated and statistically nonsignificant in the recognition task. In the source retrieval task, however, material effects in the older group were comparable in both onset latency and magnitude with those of the young subjects. Thus, the failure of older adults to demonstrate differential cue processing in tests of recognition memory likely reflects the adoption of a specific retrieval strategy rather than the incapacity to process retrieval cues in a goal-directed manner.

Considerable evidence suggests that attentional resources are necessary for the encoding of episodic memories, but the nature of the relationship between attention and neural correlates of encoding is unclear. Here we address this question using functional magnetic resonance imaging and a divided-attention paradigm in which competition for different types of attentional resources was manipulated. Fifteen volunteers were scanned while making animacy judgments to visually presented words and concurrently performing one of three tasks on auditorily presented words: male/female voice discrimination (control task), 1-back voice comparison (1-back task), or indoor/outdoor judgment (semantic task). The 1-back and semantic tasks were designed to compete for task-generic and task-specific attentional resources, respectively. Using the “remember/know” procedure, memory for the study words was assessed after 15 min. In the control condition, subsequent memory effects associated with later recollection were identified in the left dorsal inferior frontal gyrus and in the left hippocampus. These effects were differentially attenuated in the two more difficult divided-attention conditions. The effects of divided attention seem, therefore, to reflect impairments due to limitations at both task-generic and task-specific levels. Additionally, each of the two more difficult divided-attention conditions was associated with subsequent memory effects in regions distinct from those showing effects in the control condition. These findings suggest the engagement of alternative encoding processes to those engaged in the control task. The overall pattern of findings suggests that divided attention can impact later memory in different ways, and accordingly, that different attentional resources, including task-generic and task-specific resources, make distinct contributions to successful episodic encoding.

Studies employing event-related potentials (ERPs) during tests of recognition memory have reported differences in neural activity elicited by new test items according to the specific demands of the retrieval task, such as retrieving studied words versus pictures. The present study investigated whether differential processing of new items is possible when retrieval demands vary unpredictably on a trial-by-trial basis. In separate study-test phases, subjects encoded lists of intermixed words and pictures, and undertook retrieval tests with words as test items. Each test item was preceded by a task cue that signaled whether subjects were to attempt to retrieve a word or a picture from the study list. In the “blocked”condition, the targeted study material remained constant throughout the test, whereas in the “mixed”condition, the targeted material varied unpredictably across trials. New-item ERPs were more positive-going when words rather than pictures were targeted in the “blocked” condition, replicating previous findings, but this effect was absent in the “mixed”condition. The findings are consistent with the hypothesis that differential processing of retrieval cues depends upon the adoption of different task sets (“retrieval orientations” that develop over multiple trials and cannot be adjusted merely in response to an instructional cue. Unlike the new-item ERPs, ERPs elicited by the task cues in the mixed condition differed according to targeted material, but only on trials when there was a switch between target material. The implications of these findings for understanding the different retrieval strategies engaged when retrieval demands are consistent versus inconsistent are discussed.

Performing a secondary task concurrently with a study task has a detrimental effect on later memory for studied items. To investigate the mechanisms underlying this effect, the processing resources available for an incidental encoding task were varied by manipulating secondary task difficulty. fMRI data were acquired as volunteers ( n = 16) made animacy decisions to visually presented study words while concurrently performing either an easy or a hard auditory monitoring task. “Subsequent memory effects”-greater activity at study for words later remembered versus words later forgotten-were identified in the left ventral inferior frontal gyrus and the left anterior hippocampus. These effects did not vary according to whether the encoding task was performed concurrently with the easy or the hard secondary task. However, as secondary task difficulty increased, study-item activity declined and auditory-item activity increased in dorsolateral prefrontal and superior parietal regions that have been implicated in the support of executive and control functions. The findings suggest that dividing attention during encoding influences the probability of engaging the encoding operations that support later episodic memory, but does not alter the nature of the operations themselves. The findings further suggest that the probability of engaging these encoding operations depends on the level of general processing resources engaged in service of the study task.

Event-related potentials (ERPs) were used to investigate the neural processes underlying the distinctiveness heuristic— a response mode in which participants expect to remember vivid details of an experience and make recognition decisions based on this metacognitive expectation. One group of participants studied pictures and auditory words; another group studied visual and auditory words. Studied and novel items were presented at test as words only, with all novel items repeating after varying lags. ERP differences were seen between the word and picture groups for both studied and novel items. For the novel items, ERP differences were largest in frontal and central midline electrodes. In separate analyses, the picture group showed the greatest ERP differences between item types in a parietally based component from 550 to 1000 msec, whereas the word group showed the greatest differences in a frontally based component from 1000 to 2000 msec. The authors suggest that the distinctiveness heuristic is a retrieval orientation that facilitates reliance upon recollection to differentiate between item types. Although the picture group can use this heuristic and its retrieval orientation on the basis of recollection, the word group must engage additional postretrieval processes to distinguish between item types, reflecting the use of a different retrieval orientation.

Based on an event-related potential study by Rugg et al. [Dissociation of the neural correlates of implicit and explicit memory. Nature, 392, 595-598, 1998], we attempted to isolate the hemodynamic correlates of recollection, familiarity, and implicit memory within a single verbal recognition memory task using event-related fMRI. Words were randomly cued for either deep or shallow processing, and then intermixed with new words for yes/no recognition. The number of studied words was such that, whereas most were recognized (“hits”), an appreciable number of shallow-studied words were not (“misses”). Comparison of deep hits versus shallow hits at test revealed activations in regions including the left inferior parietal gyrus. Comparison of shallow hits versus shallow misses revealed activations in regions including the bilateral intraparietal sulci, the left posterior middle frontal gyrus, and the left frontopolar cortex. Comparison of hits versus correct rejections revealed a relative deactivation in an anterior left medial-temporal region (most likely the perirhinal cortex). Comparison of shallow misses versus correct rejections did not reveal response decreases in any regions expected on the basis of previous imaging studies of priming. Given these and previous data, we associate the left inferior parietal activation with recollection, the left anterior medial-temporal deactivation with familiarity, and the intraparietal and prefrontal responses with target detection. The absence of differences between shallow misses and correct rejections means that the hemodynamic correlates of implicit memory remain unclear.

ERPs elicited by correctly classified unstudied items in tests of yes/no recognition memory were used to investigate the neural correlates of retrieval cue processing. Items in Experiment 1 consisted of pictures and their corresponding names, allowing study and test material to be factorially crossed in four separate study–test cycles. The ERPs elicited by unstudied pictures and words were, in each case, more negative-going when the study material belonged to the alternative rather than the same class of items. These findings demonstrate that previously reported ERP “retrieval orientation effects” depend on differences in similarity between study and test items, and not on the form of the sought for material. In Experiments 2a and 2b, study materials were auditory words and pictures, and the test items were visual words. In both experiments, ERPs elicited by unstudied test words were more negative-going when pictures rather than auditory words were the study material. Thus, ERP retrieval orientation effects do not depend on the employment of a copy cue condition. It is proposed that the effects reflect differences in the processing necessary to maximize over lap between cue and memory representations.

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.

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.

In two experiments, words were presented in negatively or neutrally valenced sentences. At test, subjects made old/new recognition judgments to these words. In Experiment 2 only, for words judged old, subjects also indicated whether the words had been studied in a neutral or a negative context. In Experiment 1, left parietal old/new event-related brain potential (ERP) effects were larger and more sustained when elicited by words that had been studied in negative sentences, and a right frontal old/new effect was elicited by these words exclusively. In Experiment 2, the left parietal and right frontal effects elicited by old words correctly assigned to their study context were equivalent in size regardless of the nature of the context; a third ERP old/new effect, maximal over posterior scalp regions, was seen only for words from negative contexts. The findings indicate that incidental retrieval of emotional context gives rise to greater activation in neural systems supporting conscious recollection than does retrieval of nonemotional context. When contextual retrieval is intentional, recollection of emotional and non-emotional information are associated with equivalent engagement of these systems. The findings from Experiment 2 suggest that additional neural circuitry may be activated selectively by emotionally valenced episodic information.

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.

The effects on event-related potentials (ERPs) of within- and across-modality repetition of words and nonwords were investigated. In Experiment 1, subjects detected occasional animal names embedded in a series of words. AU items were equally likely to be presented auditorily or visually. Some words were repetitions, either within- or across-modality, of words presented six items previously. Visual-visual repetition evoked a sustained positive shift, which onset around 250 msec and comprised two topographically and temporally distinct components. Auditory-visual repetition modulated only the later of these two components. For auditory EMS, within- and across-modality repetition evoked effects with similar onset latencies. The within-modality effect was initially the larger, but only at posterior sites. In Experiment 2, critical items were auditory and visual nonwords, and target items were auditory words and visual pseudohomophones. Visual-visual nonword repetition effects onset around 450 msec, and demonstrated a more anterior scalp distribution than those evoked by auditory-visual repetition. Visual-auditory repetition evoked only a small, late-onsetting effect, whereas auditory-auditory repetition evoked an effect that, at parietal sites only, was almost equivalent to that from the analogous condition of Experiment 1. These findings indicate that, as indexed by ERF's, repetition effects both within- and across-modality are influenced by lexical status. Possible parallels with the effects of word and nonword repetition on behavioral variables are discussed.