陈述性记忆存储和巩固的神经机制

综述了陈述性记忆形成和巩固的神经机制,并对经典模型进行了修正和拓展。主要观点是：陈述性记忆的存储和巩固依赖于新皮层不同脑区之间建立新的联系,内侧颞叶在这一过程中起着重要作用;当陈述性记忆完全巩固后,信息的存储和提取完全依赖于新皮质;一个完整的陈述性记忆的信息被分别存储在解剖上分离的语义记忆与情节记忆存储系统内     

What Differentiates Declarative and Procedural Memories: Reply to Cohen,Poldrack, and Eichenbaum (1997)

Cohen, Poldrack, and Eichenbaum (1997; hereafter CPE) offer an account of the nature of individual items in memory and how they relate to one another. They argue that there are two separate memory systems, procedural and declarative (Cohen & Eichenbaum, 1993; Cohen & Squire, 1980). These systems differ in their neuroanatomic substrates, in their operating characteristics, and in the nature of the representations they use. CPE argue that representations in the declarative memory system are compositional, meaning that declarative representations may be composed of other declarative representations. Declarative memories are also flexible, meaning they can be accessed in contexts that differ from those in which they were encoded. Procedural memories, on the other hand, are neither compositional nor flexible. I will argue that there is not sufficient reason to argue that procedural and declarative memories have these distinct characteristics. Both procedural and declarative memories are arguably compositional, and both can appear flexible or inflexible, depending on testing conditions.     

Longitudinal study of declarative and procedural memory in primary school‐aged children

This study examined the development of declarative and procedural memory longitudinally in primary school‐aged children. At present, although there is a general consensus that age‐related improvements during this period can be found for declarative memory, there are conflicting data on the developmental trajectory of the procedural memory system. At Time 1 children aged around 5½ years were presented with measures of declarative and procedural memory. The tasks were then administered 12 months later. Performance on the declarative memory task was found to improve at a faster rate in comparison to the procedural memory task. The findings of the study support the view that multiple memory systems reach functional maturity at different points in development.     

What differentiates declarative and procedural memories: reply to Cohen, Poldrack, and Eichenbaum (1997)

CPE claim that procedural and declarative representations differ on two important dimensions: flexibility and compositionality. I have proposed that the apparent flexibility of a memory depends entirely on the transfer conditions. Any retest is, in some sense, a test of flexibility, because something has changed since the original encoding episodic. I have argued that if one changes something that does not provide support to memory performance, the memory will appear flexible, and resistant to changes in the environment. If one changes the very thing that the representation codes, the memory will appear inflexible and easily disrupted by changes in the environment. This principle is equally true for procedural and declarative memory. CPE contend that procedural representations lack compositionality. An ideal test of this claim would examine the representation of a task that is widely agreed to be procedural (e.g. that has been demonstrated to be learned normally by amnesic patients, and in the absence of awareness by neurologically intact subjects). Such experiments appear not to have been conducted, and the fact is that many tasks that are widely agreed to be procedural probably are not compositional. They appear to be, as CPE contend, biases in a processing system; it is hard to imagine how repetition priming could be compositional. Nevertheless, this is not true of all procedural memories. There is a good deal of evidence that motor behaviour is organised hierarchically and has compositionality. There is every reason to think that most if not all motor behaviour is procedural; motor behaviour might be driven by goals that are declarative, but the low-level operations that actually manipulate effectors are closed to consciousness, do not depend on the medial temporal lobe or diencephalon, and would therefore be classified as procedural. CPE framed their theory of differences between procedural and declarative memory systems as an account of the deficit in amnesic patients. They therefore predict that the learning of amnesic patients should not show flexibility or compositionality. There is already at least one study showing learning in amnesic patients that is as flexible as that of control participants (Knowlton & Squire, 1996). There are not, to my knowledge, data on whether the motor skill learning of amnesic patients shows compositionality, but one might expect that it would, given that it does in neurologically intact participants, and given that motor skill learning appears unimpaired in amnesic patients. Thus, the conception of declarative and procedural memory provided by CPE may not provide a complete account of amnesic performance. The anatomic distinction between procedural and declarative memory systems appears quite strong, and there is therefore reason to believe that there are accompanying computational differences. There does not, however, appear to be sufficient evidence to support those differences proposed by CPE.     

The Color Object Association Test (COAT): the development of a new measure of declarative memory for 18- to 36-month-old toddlers.

Few methods exist to measure declarative (explicit) memory in children during the toddler and preschool stages of development. We report the development and psychometric properties of a new measure of declarative memory for this age group, the Color Object Association Test (COAT). In pilot testing and large scale application of the test, the COAT was demonstrated to be a reliable and a valid measure of declarative memory for healthy children ages 18-36 months, living in a disadvantaged community. The test shows a linear developmental trajectory, which allows longitudinal examination of the development of declarative memory in children.     

Memory reactivation and consolidation during sleep

Do our memories remain static during sleep, or do they change? We argue here that memory change is not only a natural result of sleep cognition, but further, that such change constitutes a fundamental characteristic of declarative memories. In general, declarative memories change due to retrieval events at various times after initial learning and due to the formation and elaboration of associations with other memories, including memories formed after the initial learning episode. We propose that declarative memories change both during waking and during sleep, and that such change contributes to enhancing binding of the distinct representational components of some memories, and thus to a gradual process of cross-cortical consolidation. As a result of this special form of consolidation, declarative memories can become more cohesive and also more thoroughly integrated with other stored information. Further benefits of this memory reprocessing can include developing complex networks of interrelated memories, aligning memories with long-term strategies and goals, and generating insights based on novel combinations of memory fragments. A variety of research findings are consistent with the hypothesis that cross-cortical consolidation can progress during sleep, although further support is needed, and we suggest some potentially fruitful research directions. Determining how processing during sleep can facilitate memory storage will be an exciting focus of research in the coming years.The idea that memory storage is supported by events that take place in the brain while a person is sleeping is an idea that is only rarely acknowledged in the neuroscience community. At present, most memory research proceeds with no mention of any influence of sleep on memory. Nonetheless, this hypothesis is gaining empirical support. Research into connections between memory and sleep represents a burgeoning enterprise at the crossroads of traditional memory research and sleep research, an enterprise poised to provide novel insights into the human experience.This article presents some speculations about connections between memory and sleep. We entertain the notion that declarative memories are subject to modification during sleep, and that enduring storage of such memories is systematically influenced by neural events taking place during sleep. Although other types of memory may also be subject to change during sleep (see Maquet et al. 2003), we emphasize declarative memory here.This article also functions as an introduction to the set of papers selected for this special issue of Learning & Memory. These papers together outline portions of the current empirical basis for memory-sleep connections, including research in humans and in nonhuman animals. The findings are tantalizing, and yet there are undoubtedly major gaps in our knowledge about the functions of sleep and about how sleep may be related to memory storage. Future research on this topic is bound to grow in exciting and unpredictable ways. Here, we explore questions about declarative memory and sleep that may serve as a useful guide for such research.     

Medial temporal lobe memory in childhood: developmental transitions

The medial temporal lobes (MTL) support declarative memory and mature structurally and functionally during the postnatal years in humans. Although recent work has addressed the development of declarative memory in early childhood, less is known about continued development beyond this period of time. The purpose of this investigation was to explore MTL‐dependent memory across middle childhood. Children (6 –10 years old) and adults completed two computerized tasks, place learning (PL) and transitive inference (TI), that each examined relational memory, as well as the flexible use of relational learning. Findings suggest that the development of relational memory precedes the development of the ability to use relational knowledge flexibly in novel situations. Implications for the development of underlying brain areas and ideas for future neuroimaging investigations are discussed.     

Cognitive procedural learning in early Alzheimer's disease: impaired processes and compensatory mechanisms

Introduction. The aim of this study was to study cognitive procedural learning in early Alzheimer's disease (AD). Methods. Cognitive procedural learning was assessed using the Tower of Hanoi (TH) task. In order to take account of possible interactions between different systems during cognitive procedural learning, we also measured non‐verbal intellectual functions, working memory, and declarative memory. Results. Our results showed an apparent preservation of cognitive procedural learning in AD and a deleterious effect of the disease on verbal intelligence and declarative memory. Correlational analyses revealed a difference between AD patients and control participants in the type of processing they applied to the task. Conclusion. The non‐involvement of declarative memory would appear to be partly responsible for a slowdown in the cognitive procedural dynamics of AD patients. As the AD patients were unable to use their declarative memory, they were still in a problem‐solving mode at the end of the learning protocol and had to implement higher order cognitive processes (i.e., compensatory mechanisms) to perform the procedural task.     

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.     

Exploring memory in infancy: deferred imitation and the development of declarative memory

Imitation is an important means by which infants learn new behaviours. When infants do not have the opportunity to immediately reproduce observed actions, they may form a memory representation of the event which can guide their behaviour when a similar situation is encountered again. Imitation procedures can, therefore, provide insight into infant memory. The deferred imitation paradigm requires a modelled action to be reproduced following a delay, without prior motor practice. As such, deferred imitation procedures have been proposed to tap declarative memory abilities in non‐verbal populations such as infants. Contrary to the popular belief that infants form sparse or ill‐defined memories, deferred imitation research reveals that infants store and retrieve highly detailed memory representations. The specificity of detail encoded into the representation can, however, cause memory retrieval to fail at young ages. Developing the ability to identify event components which are central (the target stimulus) versus details which are peripheral (the exact context in which learning occurred) is therefore an important aspect of memory development. Using deferred imitation procedures to study the transition from constrained to flexible memory representations can thus facilitate our understanding of the development of declarative memory during the infancy period. Copyright © 2006 John Wiley & Sons, Ltd.     

Memory systems and the control of skilled action

ABSTRACT

In keeping with the dominant view that skills are largely automatic, the standard view of memory systems distinguishes between a representational declarative system associated with cognitive processes and a performance-based procedural system. The procedural system is thought to be largely responsible for the performance of well-learned skilled actions. Here we argue that most skills do not fully automate, which entails that the declarative system should make a substantial contribution to skilled performance. To support this view, we review evidence showing that the declarative system does indeed play a number of roles in skilled action.     

Place and the self: An autobiographical memory synthesis

In this article, I argue that the relationship between place and self can be accounted for by recent theoretical work on autobiographical memory. The link between place and self is conceptualized as a transitory mental representation that emerges as a “place of mine” (personal autobiographical experience) from a “place” (declarative knowledge). The function of “place of mine” is to guide personal memory and self-knowing consciousness of periods of our lives. I combine inquiries of memory, self, and place in a triadic relationship, a synthesis, suggesting a conceptual model for the phenomenon of place-related self as a sub-system of the self. This is formed by a causal progression from a physical place across time via emotional and cognitive bonds, components of the autobiographical information grounding the self, apportioned across declarative memory. Finally, using the methods of factor analysis and structural equation modeling, I show that the proposed model accounts for previous and new data on place-related identity.     

Childhood amnesia and distinctions between forms of memory: A comment on Wood, Brown, and Felton

Recently, Wood and his colleagues (1989) presented a case of childhood amnesia as evidence against the distinction between declarative and procedural memory that has sometimes been applied to human amnesia. Their argument was based on the observation that their patient showed some progress in school over the years, i.e., acquired some declarative knowledge, despite severely impaired day-to-day memory ability. We briefly review their case, together with a carefully studied second case of childhood amnesia not mentioned by Wood et al. Their argument is wrong in several ways. First, the utility of the declarative/procedural distinction for amnesia, or the utility of any other distinction between memory systems, depends on whether or not one kind of memory is impaired selectively, not on the severity of the impairment. In particular there is no requirement that one kind of memory be totally absent. Second, they have not provided the data necessary to support their argument; namely, data showing that the amount of declarative knowledge accumulated during years in school was better than would have been expected given the capacity for moment-to-moment or day-to-day memory. Indeed, the patient's moment-to-moment memory ability is better than represented, and the patient's progress in school was abnormally slow. Third, it is not clear that academic achievement scores provide a direct measure of declarative memory abilities (skill learning and recovery of function may also have contributed). We conclude that the evidence from childhood amnesia is fully consistent with the proposal that amnesia reflects a selective impairment in the formation of long-term declarative memory.     

Reactivation and Consolidation of Memory During Sleep

ABSTRACT— Recent research has shown compellingly that sleep supports the consolidation of declarative memories for events and facts. During consolidation, memories are stabilized against future interference and undergo qualitative changes with regard to their "explicitness" and underlying neural representation. In this article, we argue that declarative memory consolidation during sleep is based on covert reactivations of newly encoded memory traces in the hippocampus. During slow-wave sleep (SWS), the prominent slow oscillations act to synchronize the repeated reactivation of the newly encoded representations in hippocampal networks with the generation of spindle activity in the thalamus, supporting changes in neocortical networks that contribute to long-term memory storage. In this view, sleep plays an active role in the consolidation of memories, in which the neuronal reactivation of newly acquired memories is critical for the redistribution and integration of these memories into the network of pre-existing long-term memories.     

Early-life education may help bolster declarative memory in old age,especially for women

ABSTRACT

Although declarative memory declines with age, sex and education might moderate these weaknesses. We investigated effects of sex and education on nonverbal declarative (recognition) memory in 704 older adults (aged 58–98, 0–17 years of education). Items were drawings of real and made-up objects. Age negatively impacted declarative memory, though this age effect was moderated by sex and object-type: it was steeper for males than females, but only for real objects. Education was positively associated with memory, but also interacted with sex and object-type: education benefited women more than men (countering the age effects, especially for women), and remembering real more than made-up objects. The findings suggest that nonverbal memory in older adults is associated negatively with age but positively with education; both effects are modulated by sex, and by whether learning relates to preexisting or new information. The study suggests downstream benefits from education, especially for girls.     

Processing of representations in declarative and procedural working memory

The article investigates the relation between declarative and procedural working memory (WM; Oberauer, 2009). Two experiments test the assumption that representations in the two subsystems are selected for processing in analogous ways. Participants carried out a series of decisions on memorized lists of digits. For each decision, they had to select declarative and procedural representations. Regarding declarative representations, participants selected a memory set and a digit within this set as the input to each decision. With respect to the procedural representations, they selected a task set to be applied to the selected digit and a response within that task set. We independently manipulated the number of lists and the number of tasks to be switched among (one, two, or three; Experiment 1) and preparation time for a list switch (Experiment 2). For three effects commonly observed in task-switch studies, analogues in declarative WM were found: list-switch costs, mixing costs, and residual switch costs. List- and task-switch costs were underadditive, suggesting that declarative and procedural representations are selected separately and in parallel. The findings support the hypothesis of two analogous WM subsystems.     

Sleep in children enhances preferentially emotional declarative but not procedural memories

Although the consolidation of several memory systems is enhanced by sleep in adults, recent studies suggest that sleep supports declarative memory but not procedural memory in children. In the current study, the influence of sleep on emotional declarative memory (recognition task) and procedural memory (mirror tracing task) in 20 healthy children (10-13 years of age) was examined. After sleep, children showed an improvement in declarative memory. Separate analysis with respect to the emotional stimulus content revealed that sleep enhances the recognition of emotional stimuli (p > .001) rather than neutral stimuli (p = .084). In the procedural task, however, no sleep-enhanced memory improvement was observed. The results indicate that sleep in children, comparable to adults, enhances predominantly emotional declarative memory; however, in contrast to adults, it has no effect on the consolidation of procedural memory.     

Epistemic and non‐Epistemic Theories of Remembering

Contemporary memory sciences describe processes that are dynamic and constructive. This has led some philosophers to weaken the relationship between memory and epistemology; though remembering can give rise to epistemic success, it is not itself an epistemic success state. I argue that non‐epistemic (causal) theories will not do; they provide neither necessary nor sufficient conditions for remembering that p. I also argue that the shortcomings of the causal theory are epistemic in nature. Consequently, a theory of remembering must account for both its fundamentally epistemic nature and for its constructive and dynamic processes.