Content-specific source encoding in the human medial temporal lobe

Although the medial temporal lobe (MTL) is known to be essential for episodic encoding, the contributions of individual MTL subregions remain unclear. Data from recognition memory studies have provided evidence that the hippocampus supports relational encoding important for later episodic recollection, whereas the perirhinal cortex has been linked with encoding that supports later item familiarity. However, extant data also strongly implicate the perirhinal cortex in object processing and encoding, suggesting that perirhinal processes may contribute to later episodic recollection of object source details. To investigate this possibility, encoding activation in MTL subregions was analyzed on the basis of subsequent memory outcome while participants processed novel scenes paired with 1 of 6 repeating objects. Specifically, encoding activation correlating with later successful scene recognition memory was evaluated against that of source recollection for the object paired with the scene during encoding. In contrast to studies reporting a link between perirhinal cortex and item familiarity, it was found that encoding activation in the right perirhinal cortex correlates with successful recollection of the paired object. Furthermore, other MTL subregions also exhibited content-specific source encoding patterns of activation, suggesting that MTL subsequent memory effects are sensitive to stimulus category.     

The role of medial temporal lobe in item recognition and source recollection of emotional stimuli

Recent neuroimaging results suggest that distinct regions within the medial temporal lobe (MTL) may differentially support the episodic encoding of item and relational information for nonemotional stimuli (for a review, see Davachi, 2006). The present study was designed to assess whether these findings generalize to emotional stimuli. Behaviorally, we found that emotion reduced item recognition accuracy but did not reliably affect relational memory. fMRI analyses revealed that neutral and emotional words elicited distinct activation patterns within MTL regions predictive of subsequent memory. Consistent with previous findings for neutral words, hippocampal activation predicted later relational memory, whereas activation in the perirhinal cortex predicted successful item recognition. However, for emotional words, activation in the amygdala, hippocampus, and posterior parahippocampal cortex predicted item recognition only. These data suggest that MTL regions differentially support encoding of neutral and emotional stimuli.     

Episodic encoding and recognition of pictures and words: role of the human medial temporal lobes

In the present PET study, we examined brain activity related to processing of pictures and printed words in episodic memory. Our goal was to determine how the perceptual format of objects (verbal versus pictorial) is reflected in the neural organization of episodic memory for common objects. We investigated this issue in relation to encoding and recognition with a particular focus on medial temporal-lobe (MTL) structures. At encoding, participants saw pictures of objects or their written names and were asked to make semantic judgments. At recognition, participants made yes-no recognition judgments in four different conditions. In two conditions, target items were pictures of objects; these objects had originally been encoded either in picture or in word format. In two other conditions, target items were words; they also denoted objects originally encoded either as pictures or as words. Our data show that right MTL structures are differentially involved in picture processing during encoding and recognition. A posterior MTL region showed higher activation in response to the presentation of pictures than of words across all conditions. During encoding, this region may be involved in setting up a representation of the perceptual information that comprises the picture. At recognition, it may play a role in guiding retrieval processes based on the perceptual input, i.e. the retrieval cue. Another more anterior right MTL region was found to be differentially involved in recognition of objects that had been encoded as pictures, irrespective of whether the retrieval cue provided was pictorial or verbal in nature; this region may be involved in accessing stored pictorial representations. Our results suggest that left MTL structures contribute to picture processing only during encoding. Some regions in the left MTL showed an involvement in semantic encoding that was picture specific; others showed a task-specific involvement across pictures and words. Together, our results provide evidence that the involvement of some but not all MTL regions in episodic encoding and recognition is format specific.     

Modality effect in false recognition: evidence from Chinese characters

Using the Deese/Roediger‐McDermott (DRM) false memory method, Smith and Hunt ( 1998 ) first reported the modality effect on false memory and showed that false recall from DRM lists was lower following visual study than following auditory study, which led to numerous studies on the mechanism of modality effect on false memory and provided many competing explanations. In the present experiment, the authors tested the modality effect in false recognition by using a blocked presentation condition and a random presentation condition. The present experiment found a modality effect different from the results of the previous research; namely, false recognition was shown to be greater following visual study than following auditory study, especially in the blocked presentation condition rather than in the random presentation condition. The authors argued that this reversed modality effect may be due to different encoding and processing characteristics between Chinese characters and English words. Compared with English words, visual graphemes of critical lures in Chinese lists are likely to be activated and encoded in participants' minds, thus it is more difficult for participants to discriminate later inner graphemes from those items presented in visual modality. Hence visual presentation could lead to more false recognition than auditory presentation in Chinese lists. The results in the present experiment demonstrated that semantic activation occurring during the encoding and retrieve phases played an important role in modality effect in false recognition, and our findings might be explained by the activation‐monitoring account.     

Pictorial and verbal encoding in a short-term memory task

The present research attempted to manipulate the encoding modality, pictorial or verbal, of schematic faces with well-learned names by manipulating S’s expectations of the way the material was to be used. On every trial, a single name or face was presented, followed by another one; the S was asked to respond “same” if the stimuli had the same name, and “different” otherwise. The majority of second stimuli of any session was either names or faces. It was hypothesized that if S had encoded the first stimulus in the modality of the second, his judgment would be faster than if he had not appropriately encoded the first stimulus. Significantly slower reaction times were obtained to stimulus pairs where the second stimulus modality was infrequent. Further evidence that encoding of the first stimulus was in the frequent second stimulus modality comes from the finding that “different” responses were shorter when the stimuli differed on more than one attribute in the encoding (second stimulus) modality, regardless of the modality of the stimuli. Thus, evidence is presented that not only can verbal material be pictorially encoded (and vice versa), but that whether either verbal or pictorial material is verbally or pictorially encoded depends on S’s anticipation of what he is to do with the material.     

Storage of verbal associations is sufficient to activate the left medial temporal lobe

Neuroimaging studies have shown that memory encoding activates the medial temporal lobe (MTL). Many believe that these activations are related to novelty but it remains unproven which is critical - novelty detection or the rich associative encoding it triggers. We examined MTL activation during verbal associative encoding using functional magnetic resonance imaging. First, associative encoding activated left posterior MTL more than single word encoding even though novelty detection was matched, indicating not only that associative encoding activates the MTL particularly strongly, but also that activation does not require novelty detection. Moreover, it remains to be convincingly shown that novelty detection alone does produce such activation. Second, repetitive associative encoding produced less MTL activation than initial associative encoding, indicating that priming of associative information reduces MTL activation. Third, re-encoding familiar associations in a well-established way had a minimal effect on both memory and MTL activation, indicating that MTL activation reflects storage of associations, not merely their initial representation.     

Storage of verbal associations is sufficient to activate the left medial temporal lobe

Neuroimaging studies have shown that memory encoding activates the medial temporal lobe (MTL). Many believe that these activations are related to novelty but it remains unproven which is critical - novelty detection or the rich associative encoding it triggers. We examined MTL activation during verbal associative encoding using functional magnetic resonance imaging. First, associative encoding activated left posterior MTL more than single word encoding even though novelty detection was matched, indicating not only that associative encoding activates the MTL particularly strongly, but also that activation does not require novelty detection. Moreover, it remains to be convincingly shown that novelty detection alone does produce such activation. Second, repetitive associative encoding produced less MTL activation than initial associative encoding, indicating that priming of associative information reduces MTL activation. Third, re-encoding familiar associations in a well-established way had a minimal effect on both memory and MTL activation, indicating that MTL activation reflects storage of associations, not merely their initial representation.     

Cross-modality translations improve recognition by reducing false alarms

Conway and Gathercole [(1990). Writing and long-term memory: Evidence for a “translation” hypothesis. The Quarterly Journal of Experimental Psychology, 42, 513–527] proposed a translation account to explain why certain types of encoding produce benefits in memory: Switching modalities from what is presented to what is encoded enhances item distinctiveness. We investigated this hypothesis in a recognition experiment in which the presentation modality of a study list (visual vs. auditory) and the encoding activity (speaking vs. typing vs. passive encoding) were manipulated between-subjects. Manipulating encoding activity between-subjects ruled out any potential influence of the relationally distinct processing that can occur in a within-subject manipulation (in which all previous translation effects have been demonstrated). We found no overall difference in memory for words presented auditorily vs. visually nor for visual vs. auditory encoding, but critically presentation modality and encoding activity did interact. Translating from one modality to another – particularly from auditory presentation to visual encoding (typing) – led to the best memory discrimination. This was largely because of reduced false alarms, not increased hits, consistent with the distinctiveness heuristic.     

A mini-review of fMRI studies of human medial temporal lobe activity associated with recognition memory

This review considers event-related functional magnetic resonance imaging (fMRI) studies of human recognition memory that have or have not reported activations within the medial temporal lobes (MTL). For comparisons both between items at study (encoding) and between items at test (recognition), MTL activations are characterized as left/right, anterior/posterior, and hippocampus/surrounding cortex, and as a function of the stimulus material and relevance of item/source information. Though no clear pattern emerges, there are trends suggesting differences between item and source information, and verbal and spatial information, and a role for encoding processes during recognition tests. Important future directions are considered.     

A mini-review of fMRI studies of human medial temporal lobe activity associated with recognition memory.

This review considers event-related functional magnetic resonance imaging (fMRI) studies of human recognition memory that have or have not reported activations within the medial temporal lobes (MTL). For comparisons both between items at study (encoding) and between items at test (recognition), MTL activations are characterized as left/right, anterior/posterior, and hippocampus/surrounding cortex, and as a function of the stimulus material and relevance of item/source information. Though no clear pattern emerges, there are trends suggesting differences between item and source information, and verbal and spatial information, and a role for encoding processes during recognition tests. Important future directions are considered.     

Effects of dividing attention during encoding on perceptual priming of unfamiliar visual objects

According to the distractor-selection hypothesis (Mulligan, 2003), dividing attention during encoding reduces perceptual priming when responses to non-critical (i.e., distractor) stimuli are selected frequently and simultaneously with critical stimulus encoding. Because direct support for this hypothesis comes exclusively from studies using familiar word stimuli, the present study tested whether the predictions of the distractor-selection hypothesis extend to perceptual priming of unfamiliar visual objects using the possible/impossible object decision test. Consistent with the distractor-selection hypothesis, Experiments 1 and 2 found no reduction in priming when the non-critical stimuli were presented infrequently and non-synchronously with the critical target stimuli, even though explicit recognition memory was reduced. In Experiment 3, non-critical stimuli were presented frequently and simultaneously during encoding of critical stimuli; however, no decrement in priming was detected, even when encoding time was reduced. These results suggest that priming in the possible/impossible object decision test is relatively immune to reductions in central attention and that not all aspects of the distractor-selection hypothesis generalise to priming of unfamiliar visual objects. Implications for theoretical models of object decision priming are discussed.     

Imaging recollection and familiarity in the medial temporal lobe: a three-component model

The medial temporal lobe (MTL) plays a crucial role in supporting memory for events, but the functional organization of regions in the MTL remains controversial, especially regarding the extent to which different subregions support recognition based on familiarity or recollection. Here we review results from functional neuroimaging studies showing that, whereas activity in the hippocampus and posterior parahippocampal gyrus is disproportionately associated with recollection, activity in the anterior parahippocampal gyrus is disproportionately associated with familiarity. The results are consistent with the idea that the parahippocampal cortex (located in the posterior parahippocampal gyrus) supports recollection by encoding and retrieving contextual information, whereas the hippocampus supports recollection by associating item and context information. By contrast, perirhinal cortex (located in the anterior parahippocampal gyrus) supports familiarity by encoding and retrieving specific item information. We discuss the implications of a 'binding of item and context' (BIC) model for studies of recognition memory. This model argues that there is no simple mapping between MTL regions and recollection and familiarity, but rather that the involvement of MTL regions in these processes depends on the specific demands of the task and the type of information involved. We highlight several predictions for future imaging studies that follow from the BIC model.     

Arousal enhanced memory retention is eliminated following temporal lobe resection

The amygdala, situated in the anterior medial temporal lobe (MTL), is involved in the emotional enhancement of memory. The present study evaluated whether anterior MTL-resections attenuated arousal induced memory enhancement for pictures. Also, the effect of MTL-resections on response latencies at retrieval was assessed. Thirty-one patients with unilateral MTL-resections (17 left, 14 right) together with 16 controls participated in a forced choice memory task with pictorial stimuli varying in arousal. Response latencies increased with stimulus arousal in controls but not in patients. This was paralleled by attenuated recognition memory for moderately and highly arousing pictures in MTL-resectioned patients as compared to healthy controls. However, patients and controls did not differ in memory performance for non-arousing pictures. These results suggest that the MTL is necessary for arousal induced memory enhancement.     

Effects of level of processing on memory for affectively valenced words

In this study we tested whether subjects remember affectively valenced stimuli better than neutral stimuli when their attention is not focused on the affective charge of the stimuli during encoding. We tested memory for positive, negative, and neutral words in three memory tasks and using two encoding conditions (physical and semantic). Our results show that subjects remember affectively valenced stimuli better than neutral stimuli when they encode them according to their emotional content as well as to other features of the stimuli. However, although both positive and negative words are better remembered than neutral ones when semantically encoded, only positive words are better remembered than neutral ones when physically encoded. These results suggest that the affective charge of a stimulus (especially when positive) can affect retention even when a subject's attention has not focused on it during encoding.     

跨视听通道的相继错误记忆效应

本研究以12名大学生为被试采用DRM范式,考察了视觉与听觉通道在编码阶段的错误记忆ERP效应,以从更深层面认识错误记忆的内在加工机制。研究结果表明,在编码阶段,在300-500ms及500-700ms的时间窗视觉通道均存在相继错误记忆效应（DIM）,比听觉通道表现出较强的DIM效应,这不仅说明视觉通道与听觉通道在编码阶段具有不同的脑机制,而且表明DIM效应与学习项目的语义加工有着密切的关系。     

Homogeneity computation: How interitem similarity in visual short-term memory alters recognition

Visual short-term recognition memory for multiple stimuli is strongly influenced by the study items’ similarity to one another—that is, by their homogeneity. However, the mechanism responsible for this homogeneity effect has remained unclear. We evaluated competing explanations of this effect, using controlled sets of Gabor patches as study items and probe stimuli. Our results, based on recognition memory for spatial frequency, rule out the possibility that the homogeneity effect arises because similar study items are encoded and/or maintained with higher fidelity in memory than dissimilar study items are. Instead, our results support the hypothesis that the homogeneity effect reflects trial-by-trial comparisons of study items, which generate a homogeneity signal. This homogeneity signal modulates recognition performance through an adjustment of the subject’s decision criterion. Additionally, it seems the homogeneity signal is computed prior to the presentation of the probe stimulus, by evaluating the familiarity of each new stimulus with respect to the items already in memory. This suggests that recognition-like processes operate not only on the probe stimulus, but on study items as well.     

Functional Topography of the Cerebellum in Verbal Working Memory

Speech—both overt and covert—facilitates working memory by creating and refreshing motor memory traces, allowing new information to be received and processed. Neuroimaging studies suggest a functional topography within the sub-regions of the cerebellum that subserve verbal working memory. Medial regions of the anterior cerebellum support overt speech, consistent with other forms of motor execution such as finger tapping, whereas lateral portions of the superior cerebellum support speech planning and preparation (e.g., covert speech). The inferior cerebellum is active when information is maintained across a delay, but activation appears to be independent of speech, lateralized by modality of stimulus presentation, and possibly related to phonological storage processes. Motor (dorsal) and cognitive (ventral) channels of cerebellar output nuclei can be distinguished in working memory. Clinical investigations suggest that hyper-activity of cerebellum and disrupted control of inner speech may contribute to certain psychiatric symptoms.     

Configural representations in spatial working memory: modulation by perceptual segregation and voluntary attention

In what form are multiple spatial locations represented in working memory? The present study revealed that people often maintain the configural properties (interitem relationships) of visuospatial stimuli even when this information is explicitly task-irrelevant. However, the results also indicated that the voluntary allocation of selective attention prior to stimulus presentation, as well as feature-based perceptual segregation of relevant from irrelevant stimuli, can eliminate the influence of stimulus configuration on location-change detection performance. In contrast, voluntary attention cued to the relevant target location following presentation of the stimulus array failed to attenuate these influences. Thus, whereas voluntary selective attention can isolate or prevent the encoding of irrelevant stimulus locations and configural properties, people, perhaps due to limitations in attentional resources, reliably fail to isolate or suppress configural representations that have been encoded into working memory.     

Distinct roles for medial temporal lobe structures in memory for objects and their locations

The ability to learn and retain novel information depends on a system of structures in the medial temporal lobe (MTL) including the hippocampus and the surrounding entorhinal, perirhinal, and parahippocampal cortices. Damage to these structures produces profound memory deficits; however, the unique contribution to memory of each of these structures remains unclear. Here we have used functional magnetic resonance imaging (fMRI) to determine whether the perirhinal and parahippocampal cortices show differential memory-related activity. Based on the distinct patterns of cortical input to these two areas, we reasoned that these structures might show differential activity for spatial and object recognition memory. In each of 11 subjects, we found that the perirhinal cortex was active during both spatial and object memory encoding, while the anterior parahippocampal cortex was active only during spatial encoding. These data support the idea that MTL structures make distinct contributions to recognition memory performance.     

Arousal,valence, and memory for detail

A common finding in the emotion‐memory literature is that memory is enhanced for positively arousing stimuli and negatively arousing stimuli relative to neutral stimuli. We tested the notion that post‐stimulus elaboration is responsible for these effects. Post‐stimulus elaboration refers to the process of thinking about an event (after its offset) more frequently or more in‐depth. We tested the hypothesis by presenting participants with 36 slides depicting events that varied in arousal (low and high) and valence (positive and negative). The opportunity for elaboration was manipulated by requiring participants, during the inter‐slide interval, to complete addition problems, simply view the addition problems, or view a blank slide. Cued recall memory was tested for central and background details. Based on the post‐stimulus elaboration hypothesis it was expected that the greatest memory decline would occur for the central details of negatively and positively arousing slides when participants were required to complete addition problems (i.e., a distractor task × arousal × detail interaction). Contrary to the hypothesis, we found that filling the inter‐slide interval with a distractor task decreased memory for negative stimuli compared to positive stimuli. This effect was independent of arousal. We also found that arousal increased central detail memory for positive and negative stimuli and background detail memory for positive stimuli but not for negative stimuli. This interaction was explained on the basis of pre‐attentive encoding and cue utilisation. It was concluded that in order to understand the complex relationship between emotion and memory, future studies should include, as a minimum, the variables of valence, arousal, and detail.