The relationship between the right frontal old/new ERP effect and post-retrieval monitoring: specific or non-specific?

Post-retrieval processes are thought to be engaged when the outcome of an attempt to retrieve information from long-term memory must be monitored or evaluated. Previous research employing event-related potentials (ERPs) has implicated a specific ERP modulation – the ‘right frontal old/new effect’ – as a correlate of post-retrieval processing. In two experiments we examined whether the right frontal effect is specifically associated with processing of the products of an episodic retrieval attempt. During study, subjects in both experiments made one of two semantic judgments on serially presented pictures. In experiment 1, one study phase was followed by a source memory task, in which subjects responded ‘new’ to unstudied pictures and signaled the semantic judgment made on each studied picture. A separate study phase was followed by a task in which the studied items required a judgment about their semantic attributes. Robust right frontal effects were elicited by old items in both tasks, indicating that the effects are not selective for the monitoring of the content of information retrieved from episodic memory. In experiment 2, separate study phases were followed by test phases where semantic judgments were made either on old items (as in experiment 1), or on new items. Right frontal effects were elicited by whichever class of items, old or new, required the semantic judgment. Together, these findings indicate that the right frontal old/new effect reflects generic monitoring or decisional processes, rather than processing dedicated to the evaluation of the products of an episodic retrieval attempt.     

Feature binding in perceptual priming and in episodic object recognition: evidence from event-related brain potentials

It is argued that explicit remembering is based on so-called episodic tokens binding together all perceptual features of a visual object. In episodic recognition, these features are collectively reactivated. In support of this view, it has been shown that changing sensory features of a stimulus from study to test decreases subject's performance in an episodic recognition task, even though the changed features are irrelevant for the recognition judgment. On the other hand, repetition priming is unaffected by such manipulations of perceptual specificity. Implicit memory performance is therefore thought to depend on structural representations, so-called types, comprising only invariant perceptual features, but no exemplar-specific details. Event-related potentials (ERPs) in our study revealed electrophysiological evidence for the differential involvement of these perceptual memory traces in explicit and implicit memory tasks. Participants attended either a living-nonliving task or an episodic recognition task with visually presented objects. During test both groups of participants processed new objects and old objects, which were repeated either identically or in a mirror-reversed version. In the implicit task ERPs showed an occipitoparietal repetition effect, which was the same for identically repeated items and mirror reversals. In contrast, in the explicit task an early mid-frontal old/new effect for identical but not for mirror-reversed old objects was observed indicating involuntary access to perceptual information during episodic retrieval. A later portion of the old/new effect solely differentiated both types of old items from new ones.     

Brain responses of explicit and implicit memory: An event-related potential study

Implicit memory is acquired by an unintentional or unconscious learning. Recognition memory involves either automatic knowing or consciously controlled remembering. We provided an event-related potential paradigm capable of differentiating memory for the explicitly learned, implicitly learned and unstudied materials. In the explicit memory, we obtained both frontal (controlled retrieval) and parietal (recollection) old/new effects. In the implicit memory, we found persistent occipitotemporal activation (visual priming) and late attenuation in the temporoparietooccipital (repetition suppression). Event-related potential provides an insight into the dissociable mechanism of memory function that supports the dual process model with an enhanced temporal resolution on the dynamic process of both explicit perceptual learning and implicit perceptual priming.     

Inferior temporal stream for word processing with integrated mnemonic function.

The participation of the inferior temporal cortex in visual word perception and recognition raises several questions: Is there a directed processing stream proceeding anteriorly by continuous cortical processing? How fast are words processed within such an inferior temporal stream? Does this stream support implicit or explicit memory? To answer these questions, we analyzed the spatio-temporal relationship of event-related potentials, recorded directly from the inferior temporal cortex in epilepsy patients performing a continuous visual word recognition paradigm. Event-related potentials elicited an inferior temporal positivity in a strip along the left collateral sulcus. This potential exhibited a linear (r = 0.74) peak latency progression from posterior to anterior inferior temporal regions (approximately 15 cm/sec), indicating a directed, intracortical processing stream. Peak amplitudes and latencies showed reliable old/new effects with smaller amplitudes and shorter latencies for old as opposed to new words. Although the amplitude-old/new-effect occurred for all repeated words (e.g., implicit memory), the latency-old/new-effect occurred for correctly recognized old words only (e.g., explicit recognition). These results seem to dissociate two distinct mnemonic processes. The graded decrease of mean ITP peak amplitudes and latencies, however, does not allow us to exclude a single trace model as assumed for explicit recognition memory based on familiarity (Mandler [1980]: Psychol Rev 87:252-271). Regardless whether there is a dissociation between implicit and explicit memory in inferior temporal cortex or not, our findings are in accordance with an integrated inferior temporal processing stream for words that performs continuously semantic and mnemonic operations supporting both implicit and explicit memory.     

Effects of age on the neural correlates of retrieval cue processing are modulated by task demands

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.     

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.     

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.     

The posterior medial cortex is involved in visual but not in verbal memory encoding processing: an intracerebral recording study

The objective is to study the involvement of the posterior medial cortex (PMC) in encoding and retrieval by visual and auditory memory processing. Intracerebral recordings were studied in two epilepsy-surgery candidates with depth electrodes implanted in the retrosplenial cingulate, precuneus, cuneus, lingual gyrus and hippocampus. We recorded the event-related potentials (ERP) evoked by visual and auditory memory encoding–retrieval tasks. In the hippocampus, ERP were elicited in the encoding and retrieval phases in the two modalities. In the PMC, ERP were recorded in both the encoding and the retrieval visual tasks; in the auditory modality, they were recorded in the retrieval task, but not in the encoding task. In conclusion, the PMC is modality dependent in memory processing. ERP is elicited by memory retrieval, but it is not elicited by auditory encoding memory processing in the PMC. The PMC appears to be involved not only in higher-order top-down cognitive activities but also in more basic, rather than bottom-up activities.     

The electrophysiology of incidental and intentional retrieval: ERP old/new effects in lexical decision and recognition memory.

Event-related brain potentials (ERPs), recorded from a 128-sensor array were used to differentiate brain processes associated with intentional vs incidental memory retrieval. Two experiments examined ERP differences between old (studied) and new (non-studied) words and pseudowords while subjects performed either a recognition memory task or lexical decision task. Previous research has related a P600 old/new effect to the recollection of details, and the present experiments show that this effect was not amplified by intentional retrieval. The P600 effect was larger for words than pseudowords. An earlier (300 to 500 ms), frontally maximal, N400-like old/new effect ('FN400') was similar for words and pseudowords. A third, previously unidentified, mid-frontal, old/new effect was associated with only pseudoword recognition from 300 to 500 ms. Results are discussed with respect to dual-process theories of recognition memory.     

The memory that's right and the memory that's left: event-related potentials reveal hemispheric asymmetries in the encoding and retention of verbal information

We examined the nature and timecourse of hemispheric asymmetries in verbal memory by recording event-related potentials (ERPs) in a continuous recognition task. Participants made overt recognition judgments to test words presented in central vision that were either novel (new words) or had been previously presented in the left or right visual field (old words). An ERP memory effect linked to explicit retrieval revealed no asymmetries for words repeated at short and medium retention intervals, but at longer repetition lags (20-50 intervening words) this 'old/new effect' was more pronounced for words whose study presentation had been biased to the right hemisphere (RH). Additionally, a repetition effect linked to more implicit recognition processes (P2 amplitude changes) was observed at all lags for words preferentially encoded by the RH but was not observed for left hemisphere (LH)-encoded words. These results are consistent with theories that the RH encodes verbal stimuli more veridically whereas the LH encodes in a more abstract manner. The current findings provide a critical link between prior work on memory asymmetries, which has emphasized general LH advantages for verbal material, and on language comprehension, which has pointed to an important role for the RH in language processes that require the retention and integration of verbal information over long time spans.     

Comparative electrophysiological and hemodynamic measures of neural activation during memory-retrieval

The spatial and temporal characteristics of the brain processes underlying memory retrieval were studied with both event-related potentials (ERP) and positron emission tomography (PET) techniques. Subjects studied lists of 20 words and then performed episodic (old/new judgment) or semantic (living/nonliving decision) retrieval tasks on multiple four-item test lists, each lasting 10 sec. The PET and ERP measurements at test were assessed in relation to both the task (episodic vs. semantic) and the item (old vs. new or living vs. nonliving). Episodic retrieval was associated with increased blood flow in the right frontal lobe (Brodmann Area 10) and a sustained, slowly developing positive ERP shift recorded from the right frontopolar scalp. Semantic retrieval was associated with increased blood flow in the left frontal (Area 45) and temporal (Area 21) lobes but no clear ERP concomitant. The two retrieval tasks also differed from each other in the ERPs to single items in an early (300-500 ms) time window. Item-related comparisons yielded convergent results mainly if the retrieved information was relevant to the given task (e.g., old/new items during episodic retrieval and living/nonliving items during semantic retrieval). Episodically retrieved old items were associated with increased blood flow in the left medial temporal lobe and a transient increase in the amplitude of the late positive component (500-700 ms) of the ERP. Semantically retrieved living items were associated with increased blood flow in the left frontal cortex and anterior cingulate and a transient late frontal slow wave (700-1,500 ms) in the ERPs. These results indicate that the brain regions engaged in memory retrieval are active in either a sustained or transient manner. They map task-related processes to sustained and item-related processes to transient neural activity. But they also suggest that task-related factors can transiently affect early stages of item processing.     

Separating retrieval strategies from retrieval success: an event-related potential study of source memory

Event-related potentials (ERPs) were recorded while subjects performed two different source memory retrieval tasks. Each task was preceded by a study phase in which subjects heard an equal number of words spoken in a male or a female voice. A cue preceding each word indicated whether the subjects should make an active/passive (action) or a pleasant/unpleasant (liking) judgment to the word. In one retrieval task (the voice condition), subjects made a three-way distinction between new (unstudied) words, and words that had been spoken by the male or the female voice at study. In the second retrieval task (the task condition), subjects distinguished between new words, and words to which they had made an action or a liking judgment. All test words were presented visually. In keeping with previous findings, the differences between the ERPs to correct memory judgments for old and new items were characterised by two temporally and topographically dissociable modulations, with right-frontal and left-parietal maxima respectively. These 'old/new effects' displayed different sensitivities to successful retrieval of either voice or task information, providing further evidence that they index functionally dissociable processes. The direct comparison of the ERPs to correct rejections in the voice and task retrieval conditions revealed reliable differences over frontal scalp, suggesting that, irrespective of whether retrieval is successful, neural processing differs according to the source retrieval demands of the task.     

Distinguishing source memory and item memory: brain potentials at encoding and retrieval

Vivid memory for an episode generally includes memory for a central object or event plus memory for background context or source information. To assess neural differences between source and item memory, we used event-related potentials (ERPs) to monitor relevant memory processes at both encoding and retrieval. Participants fluent in Chinese studied Chinese words superimposed on a square or circular background during the study phase, followed by a 1-min delay. Then, memory was tested for both the words (items) and the corresponding background (source), or, in other blocks, tested for the words alone. ERPs to study-phase words differed as a function of whether the word was later remembered. These Dm effects in the interval from 400 to 600 ms, however, did not differ according to whether or not source was remembered. In contrast, ERPs to test-phase words showed clear old/new effects that did differ across conditions. When both item and source were remembered accurately, old/new effects emerged earlier and were larger in amplitude than when source memory was either incorrect or not queried. These results demonstrate that encoding processes indexed by ERPs may have primarily reflected encoding of the visual and semantic properties of these words, stressing item memory over source memory. Retrieval processes indexed by ERPs, in contrast, likely reflected a combination of item retrieval, source retrieval, and related processing engaged when people were remembering words seen earlier.     

Intact implicit learning in schizophrenia

OBJECTIVE: Schizophrenia impairs performance on explicit, but not implicit, memory tasks, indicating that conscious awareness at retrieval is a critical determinant of impaired memory. The authors investigated implicit learning, i.e., knowledge acquisition in the absence of conscious awareness, in patients with schizophrenia. METHOD: An artificial grammar learning task was used to assess implicit learning in 48 patients with schizophrenia and 24 healthy comparison subjects. The subjects were first presented with letter strings that were generated according to the rules of a finite-state grammar paradigm. They were then required to indicate whether new letter strings were "grammatical," depending on whether or not the strings corresponded to the rules. IQ, working memory, explicit memory, verbal fluency, and speed of processing were also assessed. RESULTS: Patients performed significantly worse than the comparison subjects on cognitive tasks that assessed episodic memory, verbal fluency, working memory, and speed of processing. In contrast, patients classified as being correct more grammatical than nongrammatical letter strings, and the magnitude of the difference was similar to that observed in healthy comparison subjects. CONCLUSIONS: Implicit learning, as assessed with an artificial grammar learning task, is intact in patients with schizophrenia. Conscious awareness might be a critical determinant of memory impairment both at encoding and at retrieval.     

Implicit and explicit memory in patients with obsessive-compulsive disorder: an event-related potential study

The implicit and explicit memory in patients with obsessive-compulsive disorder (OCD) was investigated using the event-related potential (ERP). For the assessment of implicit memory, a lexical decision task was administered. Among a total of 320 words and 140 non-words, 200 words were repeated, while the remaining 120 words and the 140 non-words were not repeated. For explicit memory, a continuous recognition task was administered, in which 280 words were repeated and 100 were not repeated. On the recognition task, both the controls and OCD patients showed more positivity to the old words than to the new words during the 200-600 ms period post-stimulus. Both groups showed faster response time to the old words than to the new words. On the lexical decision task, the controls showed the old/new effect during the 200-500 ms period post-stimulus, while the OCD patients did not. However, OCD patient showed faster response time to the old words than to the new words, although OCD patients showed prolonged response times to the old words compared to the controls. These results indicate that OCD patients have preserved explicit and implicit memory. The absence of old/new effect on ERP in OCD patients was discussed in terms of dysfunction of frontostriatal system, which plays an important role in both OCD and implicit memory.     

The control of memory retrieval: insights from event-related potentials

Effective performance on episodic retrieval tasks requires the ability to flexibly adapt to changing retrieval demands ('retrieval orientations'; [M.D. Rugg, E.L. Wilding, Retrieval processing and episodic memory, Trends Cogn. Sci. 4 (2000) 108-115]). We used event-related potentials (ERP) to examine whether maintaining a specific retrieval orientation and changing flexibly between different retrieval demands are mediated by the same brain systems or whether dissociable aspects of cognitive control are involved. Sixteen participants performed two recognition memory tasks. One required mere old/new decisions for words (general task), whereas the other task required the additional retrieval of each word's study font typeface (specific task). Furthermore, the participants either were asked to perform the same task continuously or to switch between the two tasks after every second test word. ERPs elicited by correctly rejected new (unstudied) words were analyzed. This enabled us to examine the ERP correlates of having adapted and maintained a task instruction as required during continuous blocks and of flexibly changing between retrieval demands during alternating blocks. The ERP analysis revealed more positive-going ERP slow waves for alternating blocks than for continuous blocks over bilateral frontal recording sites. This effect started around 250 ms after the test word and extended for several hundred milliseconds. As it was present for trials requiring a switch to the other task or to stay on the same task between 500 and 750 ms and no differences between the latter two trial types were obtained, it can be assumed that it is more related to general coordination requirements in alternating blocks, rather than to the actual control required to switch the retrieval task set. In addition, contrasting ERPs for the two task types revealed more positive-going ERP slow waves in the specific task than in the general task in the continuous blocks at lateral frontal recording sites between 250 and 700 ms. Together, these findings suggest that there are electrophysiologically dissociable aspects of cognitive control, namely for adapting and maintaining a retrieval orientation and for flexibly changing between varying retrieval demands.     

Electrophysiological evidence for the influence of unitization on the processes engaged during episodic retrieval: enhancing familiarity based remembering

Episodic memory depends upon multiple dissociable retrieval processes. Here we investigated the degree to which the processes engaged during successful retrieval are dependent on the properties of the representations that underlie memory for an event. Specifically we examined whether the individual elements of an event can, under some conditions, be unitized, leading to an enhancement of familiarity based responding. Retrieval processes were examined using event-related potential (ERPs) old/new effects, recorded during an associative recognition memory task. The nature of to-be-remembered information was manipulated by using word-pairs as stimuli. At study, participants were asked to remember word-pairs sharing an association (traffic-jam); association+semantic relationship (lemon-orange); or a semantic relationship only (cereal-bread). A behavioural pre-test revealed that association word-pairs were rated as having the most unitized representation. At test, participants were required to recognize if word-pairs were presented in the same pairing as at study, were rearranged from at study, or were entirely new. Behavioural recognition performance was clearly influenced by the nature of the to-be-remembered stimuli, memory being strongest for pairs related purely by association, and weakest for semantic only pairs. ERP old/new effects recorded at test also showed significant differences in the neural correlates of retrieval, depending on stimulus characteristics. The bilateral frontal old/new effect (typically associated with familiarity) was solely elicited by association only pairs. By contrast, the left parietal old/new effect (associated with recollection) was elicited equally by all three conditions. In addition, the late right frontal old/new effect (typically associated with some form of strategic/executive processing) was modulated. This latter effect was initially largest for association only pairs, and subsequently largest for semantic pairs. These findings suggest that the pattern of engagement of familiarity and recollection during successful episodic retrieval is dependent on the properties of the representations that underlie memory for an event.     

Clinical correlates of word recognition memory in obsessive-compulsive disorder: an event-related potential study

Memory disturbances found in obsessive-compulsive disorder (OCD) may partially be related to dysfunction of cortico-subcortical circuits. However, it is still unknown how OCD symptomatology is related to memory processing. To explore this question, event-related potentials (ERPs) were recorded in a continuous word-recognition paradigm in OCD patients with either severe or moderate scores on the Yale-Brown Obsessive-Compulsive Scale (Y-BOCS) (group S and group M, n=8 each) and in normal healthy controls (n=16). Typically ERPs to repeated items are characterized by more positive waveforms beginning approximately 250 ms post-stimulus. This "old/new effect" has been shown to be relevant for memory processing. The early old/new effect (ca. 300-500 ms) with a frontal distribution is proposed to be a neural correlate of familiarity-based recognition. The late old/new effect (post-500 ms) is assumed to reflect conscious memory retrieval processes. The OCD group S showed a normal early old/new effect and a reduced late old/new effect compared with group M and the control group, but no difference was found between group M and the control group. Source analyses for the late old/new effect showed statistically reduced cerebral activation in the anterior cingulate for OCD group S in contrast to the control group. Additionally, the early old/new effect in OCD group S was negatively correlated with the Y-BOCS total scores, and the late old/new effect was negatively correlated with obsession sub-scores. The severely, not moderately, ill OCD patients showed an impaired conscious recollection of the word-to-be-remembered, which suggested an impairment of working memory capacity in these patients due to a dysfunction in the frontal and cingulate cortex.     

Repetition and brain potentials when recognizing natural scenes: task and emotion differences

Repetition has long been known to facilitate memory performance, but its effects on event-related potentials (ERPs), measured as an index of recognition memory, are less well characterized. In Experiment 1, effects of both massed and distributed repetition on old–new ERPs were assessed during an immediate recognition test that followed incidental encoding of natural scenes that also varied in emotionality. Distributed repetition at encoding enhanced both memory performance and the amplitude of an old–new ERP difference over centro-parietal sensors. To assess whether these repetition effects reflect encoding or retrieval differences, the recognition task was replaced with passive viewing of old and new pictures in Experiment 2. In the absence of an explicit recognition task, ERPs were completely unaffected by repetition at encoding, and only emotional pictures prompted a modestly enhanced old–new difference. Taken together, the data suggest that repetition facilitates retrieval processes and that, in the absence of an explicit recognition task, differences in old–new ERPs are only apparent for affective cues.     

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.