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.     

Regional cerebral blood flow response to recognition memory versus semantic classification tasks

Two randomly assigned groups, N = 7 each, of neurologically normal patients were composed. One group, RM, had regional cerebral blood flow measurements during a recognition memory probe test; the other group, SC, had rCBF measurements during a semantic classification task, using the same stimulus and response features as the RM task. Thus, the groups differed only in the cognitive instruction they were executing. The RM group showed a significantly lower change from resting baseline to activation, in mean left hemisphere initial slope index, than did the SC group. The RM group, but not the SC group, showed a significant inverse correlation between occipital flow and accuracy of memory performance as indicated by d′. Correlations between age and hemispheric initial slope index, and between homologous left and right hemisphere sites, were also described. The results are considered to support an anatomical basis for the distinction between episodic and semantic memory and to suggest that occipital flow may diminish with accurate memory performance because of an upstream demand of blood in the medial temporal lobes.     

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.     

The right parahippocampal gyrus contributes to the formation and maintenance of bound information in working memory

Working memory is devoted to the temporary storage and on-line manipulation of information. Recently, an integrative system termed the episodic buffer has been proposed to integrate and hold information being entered or retrieved from episodic memory. Although the brain system supporting such an integrative buffer is still in debate, the medial temporal lobe appears to be a promising candidate for the maintenance of bound information. In the current work, binding was assessed by comparing two conditions in which participants had to retain three letters and three spatial locations presented either bound or separate. At the behavioral level, lower performance was found for bound information than for separate information. When contrasting the two conditions, activation in the right parahippocampal gyrus was greater for the encoding and maintenance of bound information. No activation was observed in the medial temporal lobe during the retrieval of bound information. Together, our results suggest that the parahippocampal gyrus may underlie the integrative and maintenance functions of the episodic buffer.     

Lateralized spatial and object memory encoding in entorhinal and perirhinal cortices

The perirhinal and entorhinal cortices are critical components of the medial temporal lobe (MTL) declarative memory system. Study of their specific functions using blood oxygenation level-dependent (BOLD) functional magnetic resonance imaging (fMRI), however, has suffered from severe magnetic susceptibility signal dropout resulting in poor temporal signal-to-noise (tSNR) and thus weak BOLD signal detectability. We have demonstrated that higher spatial resolution in the z-plane leads to improved BOLD fMRI signal quality in the anterior medial temporal lobes when using a 16-element surface coil array at 3 T (Tesla). Using this technique, the present study investigated the roles of the anterior medial temporal lobe, particularly the entorhinal and perirhinal cortices, in both object and spatial memory. Participants viewed a series of fractal images and were instructed to encode either the object's identity or location. Object and spatial recognition memory were tested after 18-sec delays. Both the perirhinal and entorhinal cortices were active during the object and spatial encoding tasks. In both regions, object encoding was biased to the left hemisphere, whereas spatial encoding was biased to the right. A similar hemispheric bias was evident for recognition memory. Recent animal studies suggest functional dissociations among regions of the entorhinal cortex for spatial vs. object processing. Our findings suggest that this process-specific distinction may be expressed in the human brain as a hemispheric division of labor.     

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.     

Cognitive neuroscience of episodic memory encoding

This paper presents a cognitive neuroscientific perspective on how human episodic memories are formed. Convergent evidence from multiple brain imaging studies using positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) suggests a role for frontal cortex in episodic memory encoding. Activity levels within frontal cortex can predict episodic memory encoding across a wide range of behavioral manipulations known to influence memory performance, such as those present during levels of processing and divided attention manipulations. Activity levels within specific frontal and medial temporal regions can even predict, on an item by item basis, whether an episodic memory is likely to form. Furthermore, separate frontal regions appear to participate in supplying code-specific information, including distinct regions which process semantic attributes of verbal information as well as right-lateralized regions which process nonverbal information. We hypothesize that activity within these multiple frontal regions provides a functional influence (input) to medical temporal regions that bind the information together into a lasting episodic memory trace.     

Functional asymmetry of human prefrontal cortex: encoding and retrieval of verbally and nonverbally coded information

There are several views about the organization of memory functions in the human prefrontal cortex. One view assumes a process-specific brain lateralization according to different memory subprocesses, that is, encoding and retrieval. An alternative view emphasizes content-specific lateralization of brain systems involved in memory processes. This study addresses this apparent inconsistency between process- and content-specific lateralization of brain activity by investigating the effects of verbal and nonverbal encoding on prefrontal activations during encoding and retrieval of environmental novel sounds using fMRI. An intentional memory task was applied in which subjects were required either to judge the sounds' loudness (nonverbal encoding task) or to indicate whether or not a sound can be verbally described (verbal encoding task). Retrieval processes were examined in a subsequent yes/no recognition test. In the study phase the right posterior dorsolateral prefrontal cortex (PFC) was activated in both tasks. During verbal encoding additional activation of the left dorsolateral PFC was obtained. Retrieval-related fMRI activity varied as a function of encoding task: For the nonverbal task we detected an activation focus in the right posterior dorsolateral PFC whereas an activation in the left dorsolateral PFC was observed for the verbal task. These findings indicate that the right dorsolateral PFC is engaged in encoding of auditory information irrespective of encoding task. The lateralization of PFC activity during retrieval was shown to depend on the availability of verbal codes, with left hemispheric involvement for verbally and right hemispheric activation for nonverbally coded information.     

Working and long-term memory deficits in schizophrenia: is there a common prefrontal mechanism?

This study tested the hypothesis that dorsolateral prefrontal cortex deficits contribute to both working memory and long-term memory disturbances in schizophrenia. It also examined whether such deficits were more severe for verbal than nonverbal stimuli. Functional magnetic resonance imaging was used to assess cortical activation during performance of verbal and nonverbal versions of a working memory task and both encoding and recognition tasks in 38 individuals with schizophrenia and 48 healthy controls. Performance of both working memory and long-term memory tasks revealed disturbed dorsolateral prefrontal cortex activation in schizophrenia, although medial temporal deficits were also present. Some evidence was found for more severe cognitive and functional deficits with verbal than nonverbal stimuli, although these results were mixed.     

Visual search enhances subsequent mnemonic search

We examined how the performance of a visual search task while studying a list of to-be-remembered words affects subsequent memory for those words by humans. Previous research had suggested that episodic context encoding is facilitated when the study phase of a memory experiment requires, or otherwise encourages, a visual search for the to-be-remembered stimuli, and theta-band oscillations are more robust when animals are searching their environment. Moreover, hippocampal theta oscillations are positively correlated with learning in animals. We assumed that a visual search task performed during the encoding of words for a subsequent memory test would induce an exploratory state that would mimic the one that is induced in animals when performing exploratory activities in their environment, and that the encoding of episodic traces would be improved as a result. The results of several experiments indicated that the performance of the search task improved free recall, but the results did not extend to yes–no or forced choice recognition memory testing. We propose that visual search tasks enhance the encoding of episodic context information but do not enhance the encoding of to-be-remembered words.     

Bi-hemispheric engagement in the retrieval of autobiographical episodes

Functional magnetic resonance imaging (fMRI) was used to study the neural correlates of neutral, stressful, negative and positive autobiographical memories. The brain activity produced by these different kinds of episodic memory did not differ significantly, but a common pattern of activation for different kinds of autobiographical memory was revealed that included (1) largely bilateral portions of the medial and superior temporal lobes, hippocampus and parahippocampus, (2) portions of the ventral, medial, superior and dorsolateral prefrontal cortex, (3) the anterior and posterior cingulate, including the retrosplenial, cortex, (4) the parietal cortex, and (5) portions of the cerebellum. The brain regions that were mainly activated constituted an interactive network of temporal and prefrontal areas associated with structures of the extended limbic system. The main bilateral activations with left-sided preponderance probably reflected reactivation of complex semantic and episodic self-related information representations that included previously experienced contexts. In conclusion, the earlier view of a strict left versus right prefrontal laterality in the retrieval of semantic as opposed to episodic autobiographical memory, may have to be modified by considering contextual variables such as task demands and subject variables. Consequently, autobiographical memory integration should be viewed as based on distributed bi-hemispheric neural networks supporting multi-modal, emotionally coloured components of personal episodes.     

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.     

Neural substrates of successful working memory and long-term memory formation in a relational spatial memory task

Working memory (WM) tasks may involve brain activation actually implicated in long-term memory (LTM). In order to disentangle these two memory systems, we employed a combined WM/LTM task, using a spatial relational (object-location) memory paradigm and analyzed which brain areas were associated with successful performance for either task using fMRI. Critically, we corrected for the performance on the respective memory task when analyzing subsequent memory effects. The WM task consisted of a delayed-match-to-sample task assessed in an MRI scanner. Each trial consisted of an indoor or outdoor scene in which the exact configuration of four objects had to be remembered. After a short delay (7–13 s), the scene was presented from a different angle and spatial recognition for two objects was tested. After scanning, participants received an unexpected subsequent recognition memory (LTM) task, where the two previously unprobed objects were tested. Brain activity during encoding, delay phase and probe phase was analyzed based on WM and LTM performance. Results showed that successful WM performance, when corrected for LTM performance, was associated with greater activation in the inferior frontal gyrus and left fusiform gyrus during the early stage of the maintenance phase. A correct decision during the WM probe was accompanied by greater activation in a wide network, including bilateral hippocampus, right superior parietal gyrus and bilateral insula. No voxels exhibited supra-threshold activity during the encoding phase, and we did not find any differential activity for correct versus incorrect trials in the WM task when comparing LTM correct versus LTM incorrect trials.     

Do humans use episodic memory to solve a <Emphasis Type="Italic">What</Emphasis>-<Emphasis Type="Italic">Where</Emphasis>-<Emphasis Type="Italic">When</Emphasis> memory task?

What-Where-When (WWW) memory tasks have been used to study episodic(-like) memory in non-human animals. In this study, we investigate whether humans use episodic memory to solve such a WWW memory task. Participants are assigned to one of two treatments, in which they hide different coin types (what) in different locations (where) on two separate occasions (when). In the Active treatment, which mimics the animal situation as closely as possible, participants are instructed to memorize the WWW information; in the Passive treatment, participants are unaware of the fact that memory will be tested. In both groups, the majority of participants report using a mental time travel strategy to solve the task, and performance on a different episodic memory test significantly predicts performance on the WWW memory task. This suggests that the WWW memory task is a good test of episodic memory in humans. Participants remember locations and coins from the first hiding session better than they do those of the second hiding session, suggesting their memories may be reinforced during the second hiding session. We also investigated how well episodic memory performance predicted performance on the three aspects of the WWW memory task separately. In the Passive treatment, episodic memory performance predicts performance on all three aspects of the WWW memory task equally. However, in the Active treatment it only predicts performance on the what component. This could imply that during active encoding a different memory system is used for where and when information than during passive encoding. Encoding of what information seems to rely on episodic memory processing in both conditions.     

Spatial and temporal episodic memory retrieval recruit dissociable functional networks in the human brain

Imaging, electrophysiological studies, and lesion work have shown that the medial temporal lobe (MTL) is important for episodic memory; however, it is unclear whether different MTL regions support the spatial, temporal, and item elements of episodic memory. In this study we used fMRI to examine retrieval performance emphasizing different aspects of episodic memory in the context of a spatial navigation paradigm. Subjects played a taxi-driver game ("yellowcab"), in which they freely searched for passengers and delivered them to specific landmark stores. Subjects then underwent fMRI scanning as they retrieved landmarks, spatial, and temporal associations from their navigational experience in three separate runs. Consistent with previous findings on item memory, perirhinal cortex activated most strongly during landmark retrieval compared with spatial or temporal source information retrieval. Both hippocampus and parahippocampal cortex activated significantly during retrieval of landmarks, spatial associations, and temporal order. We found, however, a significant dissociation between hippocampal and parahippocampal cortex activations, with spatial retrieval leading to greater parahippocampal activation compared with hippocampus and temporal order retrieval leading to greater hippocampal activation compared with parahippocampal cortex. Our results, coupled with previous findings, demonstrate that the hippocampus and parahippocampal cortex are preferentially recruited during temporal order and spatial association retrieval--key components of episodic "source" memory.     

编码与提取干扰对内隐和外显记忆的非对称性影响

以往研究表明在外显记忆中,编码与提取加工存在着非对称性,但在内隐记忆中,二者的关系并不明确,因此实验采用“学习-再认”范式,考察在编码或提取中分别附加的干扰任务对词汇判断或再认产生的影响。结果证实编码与提取干扰对内隐或外显记忆都具有非对称性的影响,但又存在着差异,即编码干扰会导致随后外显记忆成绩显著减少,而提取干扰对其影响较小,相反,编码干扰对随后内隐测验中启动效应的影响较小,但提取干扰会破坏启动效应,从而为内隐记忆和外显记忆的分离提供了进一步的证据     

Simulating episodic memory deficits in semantic dementia with the TraceLink model

Although semantic dementia is primarily characterised by deficits in semantic memory, episodic memory is also impaired. Patients show poor recall of old autobiographical and semantic memories, with better retrieval of recent experiences; they can form new memories, and normal performance on pictorial recognition memory has been demonstrated. As these abnormalities in episodic memory are virtually a mirror image of those seen in the amnesic syndromes, semantic dementia poses a challenge to extant models of remote memory and amnesia. Here, we show that one such model, TraceLink, can reproduce some of the principal findings on episodic memory in semantic dementia. A loss of nodes and connections within the trace system, which can be identified with the temporal neocortical memory storage sites implicated in semantic dementia, simulates without further assumptions the findings reported above.     

Simulating episodic memory deficits in semantic dementia with the TraceLink model

Although semantic dementia is primarily characterised by deficits in semantic memory, episodic memory is also impaired. Patients show poor recall of old autobiographical and semantic memories, with better retrieval of recent experiences; they can form new memories, and normal performance on pictorial recognition memory has been demonstrated. As these abnormalities in episodic memory are virtually a mirror image of those seen in the amnesic syndromes, semantic dementia poses a challenge to extant models of remote memory and amnesia. Here, we show that one such model, TraceLink, can reproduce some of the principal findings on episodic memory in semantic dementia. A loss of nodes and connections within the trace system, which can be identified with the temporal neocortical memory storage sites implicated in semantic dementia, simulates without further assumptions the findings reported above.     

Contributions of surface and conceptual information to recognition memory

The present experiments were designed to determine whether memory for the voice in which a word is spoken is retained in a memory system that is separate from episodic memory or, instead, whether episodic memory represents both word and voice information. These two positions were evaluated by assessing the effects of study-to-test changes in voice on recognition memory after a variety of encoding tasks that varied in processing requirements. In three experiments, the subjects studied a list of words produced by six voices. The voice in which the word was spoken during a subsequent explicit recognition test was either the same as or different from the voice used in the study phase. The results showed that word recognition was affected by changes in voice after each encoding condition and that the magnitude of the voice effect was unaffected by the type of encoding task. The results suggest that spoken words are represented in long-term memory as episodic traces that contain talker-specific perceptual information.     

相继记忆模式：展现情节记忆形成脑机制的窗口

相继记忆模式在记忆形成的脑认知成像研究领域应用广泛,已成为研究者探究大脑形成记忆时活动的主要窗口。该文在介绍相继记忆模式及记忆形成过程的基础上,分析影响相继记忆效应大小和时空分布的因素,最后讨论内侧颞叶及前额叶神经网络中相关脑区如何分工、协同支持情节记忆形成。情节记忆多维度特性导致该神经网络中有关区域表现出不同形式的相继记忆效应,因此,该文提出有效分离这些脑区在记忆形成中如何分工及交互协同关系进行更为重要