Sparse but not 'grandmother-cell' coding in the medial temporal lobe

Although a large number of neuropsychological and imaging studies have demonstrated that the medial temporal lobe (MTL) plays an important role in human memory, there are few data regarding the activity of neurons involved in this process. The MTL receives massive inputs from visual cortical areas, and evidence over the last decade has consistently shown that MTL neurons respond selectively to complex visual stimuli. Here, we focus on how the activity patterns of these cells might reflect the transformation of visual percepts into long-term memories. Given the very sparse and abstract representation of visual information by these neurons, they could in principle be considered as 'grandmother cells'. However, we give several arguments that make such an extreme interpretation unlikely.     

Overcoming interference: an fMRI investigation of pattern separation in the medial temporal lobe

The medial temporal lobe (MTL) supports the formation and retrieval of long-term declarative memories, or memories for facts and everyday events. One challenge posed for this type of memory stems from the highly overlapping nature of common episodes. Within cognitive psychology, it is widely accepted that interference between information learned at different times is a major limitation on memory. In spite of several decades of intense research in the fields of interference theory and the neurobiological underpinnings of declarative memory, there is little direct evidence bearing on how the MTL resolves this interference to form accurate memories of everyday facts and events. Computational models of MTL function have proposed a mechanism in which the MTL, specifically the hippocampus, performs pattern separation, whereby overlapping representations are made less similar. However, there is little evidence bearing on how this process is carried out in the intact human MTL. Using high-resolution fMRI, we conducted a set of experiments that taxed behavioral pattern separation by using highly similar, interfering stimuli in a modified continuous recognition task. Regions within the parahippocampal gyrus demonstrated activity consistent with a "recall to reject" strategy. In contrast and critical to performing the task, activity within the hippocampus distinguished between correctly identified true stimulus repetitions, correctly rejected presentations of similar lure stimuli, and false alarms to similar lures. These data support the computational models' assertion that the hippocampus plays a key role in pattern separation.     

Perirhinal cortical contributions to object perception

The traditional theory of the medial temporal lobe (MTL) memory system asserts that the primate MTL (hippocampus, perirhinal, entorhinal and parahippocampal cortices) is exclusively involved in consolidating declarative memories. However, several recent reports have directly challenged this dogma by arguing that MTL structures also contribute to perception. Controversy remains as many of the behavioural tasks used have confounded memory with perception. We review the evidence here and highlight new studies in humans and macaques that indicate a perceptual role for MTL in the absence of such confounds. We argue that the challenge to MTL memory system theory is substantiated and that the implications are considerable, namely that most psychologists and neuroscientists have held a fundamentally flawed view of how memory is implemented in the brain.     

The medial temporal lobe and the attributes of memory

Neuroimaging and lesion studies have seemed to converge on the idea that the hippocampus selectively supports recollection. However, these studies usually involve a comparison between strong recollection-based memories and weak familiarity-based memories. Studies that avoid confounding memory strength with recollection and familiarity almost always find that the hippocampus supports both recollection and familiarity. We argue that the functional organization of the medial temporal lobe (MTL) is unlikely to be illuminated by the psychological distinction between recollection and familiarity and will be better informed by findings from neuroanatomy and neurophysiology. These findings indicate that the different structures of the MTL process different attributes of experience. By representing the widest array of attributes, the hippocampus supports recollection-based and familiarity-based memory of multiattribute stimuli.     

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     

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.     

The eyewitness suggestibility effect and memory for source

We examined the possibility that eyewitness suggestibility reflects failures of the processes by which people normally discriminate between memories derived from different sources. To test this hypothesis, misled and control subjects were tested either with a yes/no recognition test or with a "source monitoring" test designed to orient subjects to attend to information about the sources of their memories. The results demonstrate that suggestibility effects obtained with a recognition test can be eliminated by orienting subjects toward thinking about the sources of their memories while taking the test. Our findings indicate that although misled subjects are capable of identifying the source of their memories of misleading suggestions, they nonetheless sometimes misidentify them as memories derived from the original event. The extent to which such errors reflect genuine memory confusions (produced, for example, by lax judgment criteria) or conscious misattributions (perhaps due to demand characteristics) remains to be specified.     

Patterns: Basic Units of Emotional Memory

In psychoanalysis, we attempt to engage with one another in ways that enhance understanding. We tend to valorize the verbal domains, yet much of the information we seek may be relatively inaccessible to conscious awareness, obscured by trauma or by the early age at which the information was encoded. These types of information are stored within the body as sensory memories, in which it is through the patterns of the communications that meanings are derived. Affect, most particularly, is known through its patterns of prosody and intensity. Greater appreciation of the patterned forms that underlie emotional memory can help us to better locate ourselves within this infinitely complex and fertile realm of nonverbal understandings, and to learn to communicate these understandings in constructive ways.     

What, if anything, is the medial temporal lobe, and how can the amygdala be part of it if there is no such thing?

Should the medial temporal lobe (MTL) of primates--which includes allocortical structures such as the hippocampus, neocortical structures such as the parahippocampal cortex, and nuclear structures such as the basolateral amygdala--be considered a single "thing"? According to the prevailing view, here termed the reification theory, the answer is yes. According to this theory, the MTL functions as an amalgamated entity that provides the neuronal mechanisms for declarative memory; the greater the damage to the MTL or any of its components, the greater the deleterious effects on declarative memory. A countervailing view, here called the balkanization theory, holds that the various components of the MTL process and store different kinds of information. According to this theory, damage to each part of the MTL causes a unique set of behavioral deficits-some involving memory, others involving perception, and yet others involving response selection. The empirical neuropsychological evidence favors the balkanization theory, as do some new concepts in theoretical neuroanatomy.     

System consolidation of memory during sleep

Over the past two decades, research has accumulated compelling evidence that sleep supports the formation of long-term memory. The standard two-stage memory model that has been originally elaborated for declarative memory assumes that new memories are transiently encoded into a temporary store (represented by the hippocampus in the declarative memory system) before they are gradually transferred into a long-term store (mainly represented by the neocortex), or are forgotten. Based on this model, we propose that sleep, as an offline mode of brain processing, serves the ‘active system consolidation’ of memory, i.e. the process in which newly encoded memory representations become redistributed to other neuron networks serving as long-term store. System consolidation takes place during slow-wave sleep (SWS) rather than rapid eye movement (REM) sleep. The concept of active system consolidation during sleep implicates that (a) memories are reactivated during sleep to be consolidated, (b) the consolidation process during sleep is selective inasmuch as it does not enhance every memory, and (c) memories, when transferred to the long-term store undergo qualitative changes. Experimental evidence for these three central implications is provided: It has been shown that reactivation of memories during SWS plays a causal role for consolidation, that sleep and specifically SWS consolidates preferentially memories with relevance for future plans, and that sleep produces qualitative changes in memory representations such that the extraction of explicit and conscious knowledge from implicitly learned materials is facilitated.     

Non-declarative sequence learning does not show savings in relearning

Researchers have utilized the savings in relearning paradigm in a variety of settings since Ebbinghaus developed the tool over a century ago. In spite of its widespread use, we do not yet understand what type(s) of memory are measurable by savings. Specifically, can savings measure both declarative and non-declarative memories? The lack of conscious recollection of the encoded material in some studies indicates that non-declarative memories may show savings effects, but as all studies to date have used declarative tasks, we cannot be certain. Here, we administer a non-declarative task and then measure savings in relearning the material declaratively. Our results show that while material outside of awareness may show savings effects, non-declarative sequence memory does not. These data highlight the important distinction between memory without awareness and non-declarative memory.     

Exploring the role of repetition and sensory elaboration in the imagination inflation effect

Two experiments were conducted to examine whether a misattribution of specific characteristics or a misattribution of global familiarity underlies false memories as assessed through imagination inflation. Using the paradigm developed by Goff and Roediger (1998), we found that the proportion of false memories increased with repeated imagination, replicating the imagination inflation effect. False memories developed through imagination were greatest in conditions that forced participants to include sensory detail in their imaginings. Finally, conscious recollection more often accompanied false memories in perceptually detailed imagination conditions, whereas feelings of familiarity more often accompanied false memories in conditions that lacked sensory cues. These results suggest that imagination that contains more perceptual information leads to more elaborate memory representations containing specific characteristics that can be confused with actually performed actions. Confusion based on these representations, as opposed to confusion based on processing fluency, is more likely to lead to false memories.     

Contributions of the medial temporal lobe to declarative memory retrieval: manipulating the amount of contextual retrieval

We investigated how the hippocampus and its adjacent mediotemporal structures contribute to contextual and noncontextual declarative memory retrieval by manipulating the amount of contextual information across two levels of the same contextual dimension in a source memory task. A first analysis identified medial temporal lobe (MTL) substructures mediating either contextual or noncontextual retrieval. A linearly weighted analysis elucidated which MTL substructures show a gradually increasing neural activity, depending on the amount of contextual information retrieved. A hippocampal engagement was found during both levels of source memory but not during item memory retrieval. The anterior MTL including the perirhinal cortex was only engaged during item memory retrieval by an activity decrease. Only the posterior parahippocampal cortex showed an activation increasing with the amount of contextual information retrieved. If one assumes a roughly linear relationship between the blood-oxygenation level-dependent (BOLD) signal and the associated cognitive process, our results suggest that the posterior parahippocampal cortex is involved in contextual retrieval on the basis of memory strength while the hippocampus processes representations of item-context binding. The anterior MTL including perirhinal cortex seems to be particularly engaged in familiarity-based item recognition. If one assumes departure from linearity, however, our results can also be explained by one-dimensional modulation of memory strength.     

Retrieval‐induced forgetting of emotional and unemotional autobiographical memories

This experiment extended the retrieval‐induced forgetting (RIF) procedure from simple, episodic information to emotional and unemotional autobiographical memories. In the elicitation phase, participants generated specific memories from their past in response to negative, neutral, or positive category cues. In the retrieval‐practice phase, they practised retrieving (and elaborated further on) some of the memories for some of the categories. In the final test phase, they tried to recall all memories. Memories that received retrieval practice were recalled more often on final test than baseline memories, whereas memories that were not practised, yet competed with practised memories via a shared category cue, were recalled less often than baseline memories. We discuss the roles of inhibition, competition, emotion, and self‐relevance, and consider what laboratory manipulations of memory might reveal about everyday and pathological personal memory.     

Auditory memory can be object based

Identifying how memories are organized remains a fundamental issue in psychology. Previous work has shown that visual short-term memory is organized according to the object of origin, with participants being better at retrieving multiple pieces of information from the same object than from different objects. However, it is not yet clear whether similar memory structures are employed for other modalities, such as audition. Under analogous conditions in the auditory domain, we found that short-term memories for sound can also be organized according to object, with a same-object advantage being demonstrated for the retrieval of information in an auditory scene defined by two complex sounds overlapping in both space and time. Our results provide support for the notion of an auditory object, in addition to the continued identification of similar processing constraints across visual and auditory domains. The identification of modality-independent organizational principles of memory, such as object-based coding, suggests possible mechanisms by which the human processing system remembers multimodal experiences.     

Neural correlates underlying true and false associative memories

Despite the fact that associative memory studies produce a large number of false memories, neuroimaging analyses utilizing this paradigm typically focus only on neural activity mediating successful retrieval. The current study sought to expand on this prior research by examining the neural basis of both true and false associative memories. Though associative false memories are substantially different than those found in semantic or perceptual false memory paradigms, results suggest that associative false memories are mediated by similar neural mechanisms. Specifically, we found increased frontal activity that likely represents enhanced monitoring and evaluation compared to that needed for true memories and correct rejections. Results also indicated that true, and not false associative memories, are mediated by neural activity in the MTL, specifically the hippocampus. Finally, while activity in early visual cortex distinguished true from false memories, a lack of neural differences between hits and correct rejections failed to support previous findings suggesting that activity in early visual cortex represents sensory reactivation of encoding-related processing.     

Cognitive processes supporting episodic memory formation in childhood: The role of source memory,binding, and executive functioning

Episodic memories contain various forms of contextual detail (e.g., perceptual, emotional, cognitive details) that need to become integrated. Each of these contextual features can be used to attribute a memory episode to its source, or origin of information. Memory for source information is one critical component in the formation of episodic memories. Likewise, the establishment of episodic memories also requires binding, which reflects the process of encoding the relations among stimuli and provides the experience that certain features of a memory episode belong together. The aims of the present review are to explore the roles of (1) cognitive and neural mechanisms involved in source memory and binding and how the development of these cognitive processes relates to episodic memory formation in childhood and (2) other higher-order cognitive processes, namely executive functioning, in early episodic memory development. We conclude by examining the challenges within this field of research, highlighting the role of other cognitive processes (e.g., sense of self, language, use of strategies) that may contribute to episodic memory formation, addressing areas that can be improved with additional research, and exploring directions for future work.     

Face your fears: attenuating remote fear memories by reconsolidation-updating

Traumatic events generate some of the most enduring memories, yet little is known about how long-lasting fear memories can be attenuated. In this review, we collect the surprisingly sparse evidence on remote fear memory attenuation from both animal and human research. What is becoming apparent is twofold: although remote fear memories are more resistant to change compared with recent ones, they can nevertheless be attenuated when interventions are targeted toward the period of memory malleability instigated by memory recall, the reconsolidation window. We describe the physiological mechanisms underlying remote reconsolidation-updating approaches and highlight how they can be enhanced through interventions promoting synaptic plasticity. By capitalizing on an intrinsically relevant phase of memory, reconsolidation-updating harbors the potential to permanently alter remote fear memories.     

Out of one's mind: a study of involuntary semantic memories

The study of memories that pop into one's mind without any conscious attempt to retrieve them began only recently. While there are some studies on involuntary autobiographical memories (e.g., ) research on involuntary semantic memories or mind-popping is virtually non-existent. The latter is defined as an involuntary conscious occurrence of brief items of one's network of semantic knowledge. The recall of these items (e.g., a word, a name, a tune) is not accompanied by additional contextual information and/or involvement of self-a standard feature of involuntary autobiographical memories. The paper reports several diary and questionnaire studies which looked into the nature and frequency of occurrence of these memories. The data show that people do experience involuntary semantic memories which tend to occur without any apparent cues while being engaged in relatively automatic activities. Possible mechanisms of involuntary semantic memories are discussed (e.g., very long-term priming), and the results of the study provide information on the possible duration of the priming effects in everyday life. Related theoretical and methodological issues and future avenues of research in this neglected area are outlined.