“Wanted!” The effects of reward on face recognition: electrophysiological correlates

The aim of the present study was to uncover the temporal dynamics of face recognition as a function of reward. Event-related potentials (ERPs) were recorded during the encoding and the subsequent old/new memory test in response to faces that could be associated with a monetary reward. The behavioral results showed that faces associated with reward at both encoding and retrieval were recognized better than the unrewarded ones. ERP responses highlighted that successful encoding predictive of subsequent memory was greater for faces associated with reward than for faces without reward-driven motivational learning. At retrieval, an early positive-going component was elicited for potentially rewarded faces on frontal regions, while the occipito-temporal N170 component showed priming effects as a function of reward. Later on, larger centro-parietal ERP components, related to recognition memory, were found selectively for reward-associated faces. Remarkably, electrophysiological responses varied in a graded manner, with the largest amplitude yielded by faces with double reward, followed by faces associated with reward only at encoding. Taken together, the present data show that the processing of outcome expectations affects face structural encoding and increases memory efficiency, yielding a robust and sustained modulation over frontal and temporal areas where reward and memory mechanisms operate in conjunction.     

Regional and inter-regional theta oscillation during episodic novelty processing

Recent event-related potential (ERP) and functional magnetic resonance imaging (fMRI) studies suggest that novelty processing may be involved in processes that recognize the meaning of a novel sound, during which widespread cortical regions including the right prefrontal cortex are engaged. However, it remains unclear how those cortical regions are functionally integrated during novelty processing. Because theta oscillation has been assumed to have a crucial role in memory operations, we examined local and inter-regional neural synchrony of theta band activity during novelty processing. Fifteen right-handed healthy university students participated in this study. Subjects performed an auditory novelty oddball task that consisted of the random sequence of three types of stimuli such as a target (1000 Hz pure tone), novel (familiar environmental sounds such as dog bark, buzz, car crashing sound and so on), and standard sounds (950 Hz pure tone). Event-related spectra perturbation (ERSP) and the phase-locking value (PLV) were measured from human scalp EEG during task. Non-parametric statistical tests were applied to test for significant differences between stimulus novelty and stimulus targets in ERSP and PLV. The novelty P3 showed significant higher amplitude and shorter latency compared with target P3 in frontocentral regions. Overall, theta activity was significantly higher in the novel stimuli compared with the target stimuli. Specifically, the difference in theta power between novel and target stimuli was most significant in the right frontal region. This right frontal theta activity was accompanied by phase synchronization with the left temporal region. Our results imply that theta phase synchronization between right frontal and left temporal regions underlie the retrieval of memory traces for unexpected but familiar sounds from long term memory in addition to working memory retrieval or novelty encoding.     

Interrupted actions affect output monitoring and event-related potentials (ERPs)

Memory for performed and interrupted actions was measured on source recognition and source recall tests in order to investigate output monitoring (i.e., memory for actions). Event-related potentials (ERPs) were recorded during the source recognition test to provide insight into the neural basis of output monitoring (OM). Source identification and recall of performed actions was greater than interrupted actions, thereby replicating the enactment effect. Examination of memory errors revealed that interrupted actions were more often mistaken as performed actions. The ERP data indicated that brain activity elicited by performed actions differed from interrupted and new actions. A clear difference in temporal onset of two ERP effects (i.e., a central-parietal and a frontal ERP difference) was observed, and it supports the previous hypothesis that two distinct processes support OM and source monitoring judgements. The pattern of frontal ERP differences suggested that interrupted actions prompted people to use more systematic decision processes overall to make OM judgements. Central-parietal ERP effects suggested that sensori-motor information was not recollected for interrupted actions--rather OM judgements were based on cognitive operations in this case.     

Interrupted actions affect output monitoring and event-related potentials (ERPs)

Memory for performed and interrupted actions was measured on source recognition and source recall tests in order to investigate output monitoring (i.e., memory for actions). Event-related potentials (ERPs) were recorded during the source recognition test to provide insight into the neural basis of output monitoring (OM). Source identification and recall of performed actions was greater than interrupted actions, thereby replicating the enactment effect. Examination of memory errors revealed that interrupted actions were more often mistaken as performed actions. The ERP data indicated that brain activity elicited by performed actions differed from interrupted and new actions. A clear difference in temporal onset of two ERP effects (i.e., a central-parietal and a frontal ERP difference) was observed, and it supports the previous hypothesis that two distinct processes support OM and source monitoring judgements. The pattern of frontal ERP differences suggested that interrupted actions prompted people to use more systematic decision processes overall to make OM judgements. Central-parietal ERP effects suggested that sensori-motor information was not recollected for interrupted actions—rather OM judgements were based on cognitive operations in this case.     

Fearful expressions enhance recognition memory: electrophysiological evidence

Facial expressions play a key role in affective and social behavior. However, the temporal dynamics of the brain responses to emotional faces remain still unclear, in particular an open question is at what stage of face processing expressions might influence encoding and recognition memory. To try and answer this question we recorded the event-related potentials (ERPs) elicited in an old/new recognition task. A novel aspect of the present design was that whereas faces were presented during the study phase with either a happy, fearful or neutral expression, they were always neutral during the memory retrieval task. The ERP results showed three main findings: An enhanced early fronto-central positivity for faces encoded as fearful, both during the study and the retrieval phase. During encoding subsequent memory (Dm effect) was influenced by emotion. At retrieval the early components P100 and N170 were modulated by the emotional expression of the face at the encoding phase. Finally, the later ERP components related to recognition memory were modulated by the previously encoded facial expressions. Overall, these results suggest that face recognition is modulated by top-down influences from brain areas associated with emotional memory, enhancing encoding and retrieval in particular for fearful emotional expressions.     

Recall and stem-completion priming have different electrophysiological correlates and are modified differentially by directed forgetting

The notion that different aspects of memory are assessed by explicit and implicit memory tests was supported by behavioral and electrophysiological results. In a study-test procedure, 24 subjects were instructed to remember some words and to forget other words. Free recall and cued recall were better for words associated with the remember instruction, whereas directed forgetting did not influence stem completion (an implicit memory test). Event-related brain potentials elicited during study differed as a function of subsequent memory performance for free recall and cued recall, but not for stem completion. These results implicate encoding differences in the distinction between the 2 types of memory test. Factors governing whether explicit retrieval affects performance on an implicit memory test, mechanisms that may underlie directed-forgetting effects, and the importance of electrophysiological correlates of memory are also discussed.     

The effect of specific test queries on source-monitoring event-related potentials

Source-monitoring decision processes were manipulated during retrieval while event-related potentials (ERPs) were recorded. Words were either seen or heard at study, and memory for modality was measured on two separate yes-no source tests. Decision processes were varied across the two tests by asking participants to respond to leading questions. One leading question asked if the items were seen at study, whereas the second question asked if the items were heard at study (cf., Marsh & Hicks, 1998). Behavioral responses indicated that leading questions altered the way in which memory was evaluated to determine the source of information. Varying the decision processes affected frontal--but not parietal ERPs--indicating that frontal ERPs reflect processing that is used to evaluate activated information. Furthermore, left and right frontal ERP activity was affected by the combination of test query and type of source supporting the hypothesis that both the right and left frontal lobes contribute to memory retrieval processes. The pattern of frontal ERP effects supports the hypothesis that activation in right frontal areas reflect basic decision processes that are used to determine source and that the left frontal lobes are recruited when more systematic processing is required by the test context (cf., Nolde, Johnson, & Raye, 1998b).     

Electrophysiological correlates of encoding and retrieving emotional events

This study examined the impact of emotional content on encoding and retrieval processes. Event-related potentials were recorded in a source recognition memory task. During encoding, a posterior positivity for positive and negative pictures (250-450 ms) that presumably reflects attentional capturing of emotionally valenced stimuli was found. Additionally, positive events, which were also rated as less arousing than negative events, gave rise to anterior and posterior slow wave activity as compared with neutral and negative events and also showed enhanced recognition memory. It is assumed that positive low-arousing events enter controlled and elaborated encoding processes that are beneficial for recognition memory performance. The high arousal of negative events may interfere with controlled encoding mechanisms and attenuate item recognition and the quality of remembering. Moreover, topographically distinct late posterior negativities were obtained for the retrieval of the context features location and time that support the view that this component reflects processes in service of reconstructing the study episode by binding together contextual details with an item and that varies with the kind of episodic detail to be retrieved.     

Monetary rewards influence retrieval orientations

Reward anticipation during learning is known to support memory formation, but its role in retrieval processes is so far unclear. Retrieval orientations, as a reflection of controlled retrieval processing, are one aspect of retrieval that might be modulated by reward. These processes can be measured using the event-related potentials (ERPs) elicited by retrieval cues from tasks with different retrieval requirements, such as via changes in the class of targeted memory information. To determine whether retrieval orientations of this kind are modulated by reward during learning, we investigated the effects of high and low reward expectancy on the ERP correlates of retrieval orientation in two separate experiments. The reward manipulation at study in Experiment 1 was associated with later memory performance, whereas in Experiment 2, reward was directly linked to accuracy in the study task. In both studies, the participants encoded mixed lists of pictures and words preceded by high- or low-reward cues. After 24?h, they performed a recognition memory exclusion task, with words as the test items. In addition to a previously reported material-specific effect of retrieval orientation, a frontally distributed, reward-associated retrieval orientation effect was found in both experiments. These findings suggest that reward motivation during learning leads to the adoption of a reward-associated retrieval orientation to support the retrieval of highly motivational information. Thus, ERP retrieval orientation effects not only reflect retrieval processes related to the sought-for materials, but also relate to the reward conditions with which items were combined during encoding.     

Adapting to changing memory retrieval demands: Evidence from event-related potentials

This study investigated preparatory processes involved in adapting to changing episodic memory retrieval demands. Event-related potentials (ERPs) were recorded while participants performed a general old/new recognition task and a specific task that also required retrieval of perceptual details. The relevant task remained either constant or changed (predictably or randomly) across trials. Responses were slowed when participants switched from the specific to the general task but not vice versa. Hence, asymmetrical switch costs were observed, suggesting that retrieval preparation is dependent not only on the current retrieval goal but also influenced by recent retrieval attempts. Consistently, over posterior scalp regions ERPs associated with advance preparation were modulated by the preceding task, reflecting increased attentional selection requirements for the general task, and by the foreknowledge about the task sequence. When retrieval demands remained constant, frontal slow-waves elicited by retrieval-cues were more positive going for the specific task, indicating full implementation of a retrieval orientation that allows more efficient retrieval of perceptual details.     

Neural correlates of the formation and realization of delayed intentions

Prospective memory (PM) can be thought of as the ability to successfully form and later realize intentions that must be delayed over some period of time. In this study, event-related brain potentials were used to explore the neural activity associated with the formation and realization of an intention. Greater negativity over the frontal-polar region was associated with intention formation trials in which the intention was later realized. On PM cue trials, an N300 was associated with the detection of a cue. For PM cue trials, a late positive complex was observed that might have reflected the retrieval of an intention from memory, and a frontal slow wave was observed that might have reflected the activity of a neural system that supported disengagement from the ongoing activity when the cue was detected.     

In Search of the Self: A Positron Emission Tomography Study

Abstract—Previous work using positron emission tomography (PET) has shown that memory encoding processes are associated with preferential activation of left frontal regions of the brain, whereas retrieval processes are associated predominantly with right frontal activations. One possible reason for the asymmetry is that episodic retrieval necessarily involves reference to the self, and the self-concept may be represented (at least partially) in right frontal regions. Accordingly, the present study investigated the possibility that encoding of self-related material might also activate right frontal areas. Eight right-handed volunteers judged trait adjectives under four separate PET scan conditions: (a) relevance to self, (b) relevance to a well-known public figure, (c) social desirability, and (d) number of syllables. The results showed that self-related encoding yielded left frontal activations similar to those associated with other types of semantic encoding, but also specific activations in the right frontal lobe. It is concluded that the concept of self involves both general schematic structures and further specific components involved in episodic memory retrieval.     

Retrieval of concrete words involves more contextual information than abstract words: multiple components for the concreteness effect

The current study used the directed forgetting paradigm in implicit and explicit memory to investigate the concreteness effect. Event-related potentials (ERPs) were recorded to explore the neural basis of this phenomenon. The behavioral results showed a clear concreteness effect in both implicit and explicit memory tests; participants responded significantly faster to concrete words than to abstract words. The ERP results revealed a concreteness effect (N400) in both the encoding and retrieval phases. In addition, behavioral and ERP results showed an interaction between word concreteness and memory instruction (to-be-forgotten vs. to-be-remembered) in the late epoch of the explicit retrieval phase, revealing a significant concreteness effect only under the to-be-remembered instruction condition. This concreteness effect was realized as an increased P600-like component in response to concrete words relative to abstract words, likely reflecting retrieval of contextual details. The time course of the concreteness effect suggests advantages of concrete words over abstract words due to greater contextual information.     

Neurophysiology of successful encoding and retrieval of source memory

Event-related potentials were recorded during encoding, to identify whether brain activity predicts subsequent retrieval of spatial source information, and during retrieval, to investigate the neural correlates of successful and unsuccessful spatial context recollection. The amplitude registered during encoding for study items that were later recognized and assigned a correct source judgment was more positive than the amplitude for recognized items given incorrect source judgments; this difference started 480 msec poststimulus, predominantly at central and anterior sites. During retrieval, the waveform was more positive from 250 to 1,600 msec poststimulus when the brain had retrieved episodic information successfully than when it had failed. These findings indicate that brain electrical activity recorded during the first presentation of an item within its context predicts the subsequent retrieval of the specific spatial context. During retrieval, brain activity differed quantitatively at anterior sites and qualitatively at posterior sites according to the accuracy of source memory.     

Exploring the influence of encoding format on subsequent memory

Distinctive encoding is greatly influenced by gist-based processes and has been shown to suffer when highly similar items are presented in close succession. Thus, elucidating the mechanisms underlying how presentation format affects gist processing is essential in determining the factors that influence these encoding processes. The current study utilised multivariate partial least squares (PLS) analysis to identify encoding networks directly associated with retrieval performance in a blocked and intermixed presentation condition. Subsequent memory analysis for successfully encoded items indicated no significant differences between reaction time and retrieval performance and presentation format. Despite no significant behavioural differences, behaviour PLS revealed differences in brain–behaviour correlations and mean condition activity in brain regions associated with gist-based vs. distinctive encoding. Specifically, the intermixed format encouraged more distinctive encoding, showing increased activation of regions associated with strategy use and visual processing (e.g., frontal and visual cortices, respectively). Alternatively, the blocked format exhibited increased gist-based processes, accompanied by increased activity in the right inferior frontal gyrus. Together, results suggest that the sequence that information is presented during encoding affects the degree to which distinctive encoding is engaged. These findings extend our understanding of the Fuzzy Trace Theory and the role of presentation format on encoding processes.     

Understanding age-related reductions in visual working memory capacity: Examining the stages of change detection

Visual working memory (VWM) capacity is reduced in older adults. Research has shown age-related impairments to VWM encoding, but aging is likely to affect multiple stages of VWM. In the present study, we recorded the event-related potentials (ERPs) of younger and older adults during VWM maintenance and retrieval. We measured encoding-stage processing with the P1 component, maintenance-stage processing with the contralateral delay activity (CDA), and retrieval-stage processing by comparing the activity for old and new items (old–new effect). Older adults showed lower behavioral capacity estimates (K) than did younger adults, but surprisingly, their P1 components and CDAs were comparable to those of younger adults. This remarkable dissociation between neural activity and behavior in the older adults indicated that the P1 and CDA did not accurately assess their VWM capacity. However, the neural activity evoked during VWM retrieval yielded results that helped clarify the age-related differences. During retrieval, younger adults showed early old–new effects in frontal and occipital areas and a late central–parietal old–new effect, whereas older adults showed a late right-lateralized parietal old–new effect. The younger adults’ early old–new effects strongly resembled an index of perceptual fluency, suggesting that perceptual implicit memory was activated. The activation of implicit memory could have facilitated the younger adults’ behavior, and the lack of these early effects in older adults may suggest that they have much lower-resolution memory than do younger adults. From these data, we speculated that younger and older adults store the same number of items in VWM, but that younger adults store a higher-resolution representation than do older adults.     

Neural evidence for the contribution of holistic processing but not attention allocation to the other-race effect on face memory

Multiple mechanisms have been suggested to contribute to the other-race effect on face memory, the phenomenon of better memory performance for own-race than other-race faces. Here, two of these mechanisms, increased attention allocation and greater holistic processing during memory encoding for own-race than other-race faces, were tested in two separate experiments. In these experiments event-related potentials were measured during study (the difference due to memory, Dm) and test phase (old/new effects) to examine brain activation related to memory encoding and retrieval, allowing for selective investigations of these memory sub-processes. In Experiment 1, participants studied own-race (Caucasian) and other-race (Chinese) faces under focused or divided attention. In Experiment 2, participants studied own-race (Caucasian) and other-race (African American) faces presented upright or upside down (i.e., inverted). Both experiments showed decreases in memory performance when attention allocation or holistic processing was reduced, but these effects were similar for own-race and other-race faces. Manipulations of holistic processing, but not attention allocation, influenced the neural other-race effects during memory encoding. Inverted own-race faces showed similar neural patterns as upright other-race faces, indicating that when holistic processing of own-race faces was reduced, these faces were encoded similarly as upright other-race faces. No influences of the experimental manipulations on other-race effects during memory retrieval were found. The present study provides the first neural evidence that increased holistic processing during memory encoding contributes to the other-race effect on face memory.     

The effect of retrieval enactment on recall of subject-performed tasks and verbal tasks

The effect of retrieval enactment on memory for nouns (objects) or verbal phrases describing simple actions (e.g., “lift the box”) was addressed in two experiments. In Experiment 1, the type of object involved in the actions was manipulated, with three different types of object being used (body parts, laboratory-related objects, and external objects). In Experiment 2, the integration between the verb-noun pairs was manipulated (well-integrated vs. poorly integrated). Results from both experiments showed that whereas encoding enactment (motor encoding and verbal test) substantially improved the memory performance compared with a verbal condition (verbal encoding and verbal test), retrieval enactment (verbal encoding and motor test) had no major impact on the number of recalled nouns or phrases. Moreover, there was no additional effect of dual enactment (motor encoding and motor test). The overall pattern of the results suggests that there is a fundamental difference between motor processing at encoding and motor processing at retrieval, and the lack of encoding specificity advantage for the motor modality contradicts the view that encoding enactment of verbal commands results in storage of motor representations.     

The Compensatory Role of Implementation Intentions for Young Adults with Low Working Memory Capacity

Many factors improve prospective memory performance both inside and outside of the laboratory, including the detailed planning of the situational cue and intended action (i.e., implementation intentions). In the current study, we obtained measures of working memory capacity and laboratory event‐based prospective memory performance in college‐aged adults. Half of our participants formed an implementation intention in the prospective memory task. Because of evidence that implementation intentions increase the encoding/retrieval efficiency of the prospective memory, it was predicted that forming an implementation intention would serve as a compensatory strategy for those with low working memory ability. Our results supported this hypothesis in that working memory capacity no longer correlated with prospective memory performance when participants employed an implementation intention encoding strategy. These findings suggest that implementation intentions may be an effective way for individuals with low working memory capacity to improve their performance in an attentionally demanding prospective memory task. Copyright © 2015 John Wiley & Sons, Ltd.     

Relational information processing of novel unrelated actions by infants

Declarative memory in infants is often assessed via deferred imitation. Not much is known about the information processing basis of the memory effect found in these experiments. While in the typical deferred imitation study the order of actions remains the same during demonstration and retrieval, in two experiments with n=30 respective n=25, 10- and 11-month-old infants, the order of novel unrelated actions in demonstration and retrieval was varied (same, reversed, mixed). This allowed a separation of item-specific from item-relational information processing. In both experiments best memory performance was found when the order of target actions remained the same during encoding and recall, demonstrating that infants seem to rely on item-specific as well as item-relational information which has to be ad hoc constructed while encoding.