Memory consolidation

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on motor learning have noted that while consolidation occurs over a period of 4–6 hours during sleep, this is also true during waking hours, which may negate any role of sleep in learning. In this sense sleep would serve no special purpose to enhance consolidation of memories because it occurs independently of sleep. Other studies have examined the process of replay which has been described as a reactivation of patterns that were stimulated during a learning phase. Replay has been demonstrated in the hippocampus and this has lent support to the notion that it serves a consolidation purpose. However, replay is not specific to sleep and both rats and primates show signs during restful-awake periods. Also, replay may simply be residual activation in areas that were involved previously in the learning phase and may have no actual effect on consolidation. This reactivation of the memory traces has also been seen in non-REM sleep specifically for hippocampus-dependant memories. Researchers have noted strong reactivation of the hippocampus during sleep immediately after a learning task. This reactivation led to enhanced performance on the learned task. One such experiment had participants learn word pair associations (declarative memories) before either retention periods of sleep or periods of wakefulness. Researchers found that retrieval expectancy played a role as to whether participants were able to retain the information, as the participants that had been told about the delayed retrieval test performed better. However, their research showed that sleep was more likely to benefit the consolidation of memories if the information was relevant to future events or behaviors. Researchers following this line of work have come to assume that

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storage of episodic memories, can be established in structures apart from the hippocampal system such as the neo-cortex in the process of consolidation. Hence, while proper hippocampal functioning is necessary for the retention and retrieval of episodic memories, it is less necessary during the encoding and use of semantic memories. As memories age there are long-term interactions between the hippocampus and neo-cortex and this leads to the establishment of aspects of memory within structures aside from the hippocampus. MTT thus states that both episodic and semantic memories rely on the hippocampus and the latter becomes somewhat independent of the hippocampus during consolidation. An important distinction between MTT and the standard model is that the standard model proposes that all memories become independent of the hippocampus after several years. However, Nadel and Moscovitch have shown that the hippocampus was involved in memory recall for all remote

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course of four days in one room, and the last was taught the class over the course of four days in different rooms. The subjects were tested five days later in a completely new setting. The results of the experiment were that taking the class over a period of four days was much more effective than taking it in one single mass. Interestingly, the group that took the course over four days and in different rooms performed the best in the final retention test out of all the groups." This shows that spacing out study sessions and studying in different environments helps with retention as it provides time for the brain to consolidate the information without being interrupted by new information. The benefits of spacing were also demonstrated in an earlier study by Reder and Anderson (1982) which yielded similar results, confirming the spacing effect’s relevance and effects on learning.

592:(BLA) is involved in the encoding of significant experiences and has been directly linked to memorable events. Extensive evidence suggests that stress hormones such as epinephrine play a critical role in consolidating new memories and this is why stressful memories are recalled vividly. Studies by Gold and van Buskirk provided initial evidence for this relationship when they showed that injections of epinephrine into subjects following a training period resulted in greater long-term retention of task related memories. This study also provided evidence that the level of epinephrine injected was related to the level of retention suggesting that the level of stress or emotionality of the memory plays a role on the level of retention. It is suggested that epinephrine affects memory consolidation by activating the amygdala and studies have shown that 576:

on tasks and exhibit learning without the subject being aware that the training had ever taken place. This introduces a dissociation between the two forms of memory and the fact that one form can exist absent the other suggests separate mechanisms are involved in consolidation. Squire has proposed the procedural knowledge is consolidated in some cases by the extrapyramidal motor system. Squire demonstrated that intact learning of certain motor, perceptual, and cognitive skills can be retained in patients with amnesia. They also retain the ability to be influenced by priming effects without the patients being able to consciously recall any training session occurring.

837:). Following the same method that Nader and his associates used, Brunet induced anxiety responses in the patients by having them listen to a 30 second recording describing the circumstances of their traumatic experiences. The patients were shortly thereafter injected with propranolol, a drug that blocks stress hormone receptors in the amygdala which is implicated in neurologically representing the emotional content of memories. These patients experienced a significant reduction in PTSD symptoms months after treatment. These findings were confirmed in later studies done in 2009 by Kindt and colleagues and in 2010 by Schiller and colleagues. 550:

hippocampal damage show traces of memories and this has been used as support for the standard model because it suggests that memories are retained apart from the hippocampal system. Nadel and Moscovitch argue that these retained memories have lost the richness of experience and exist as depersonalized events that have been semanticized over time. They suggest that this instead provides support for their notion that episodic memories rely significantly on the hippocampal system but semantic memories can be established elsewhere in the brain and survive hippocampal damage.

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shock. Groups of rats were then injected with anisomycin, an antibiotic that restricts protein synthesis, at different points in time. The rats that were injected with anisomycin after consolidation had taken place, retained the fear reaction to the tone. However, the rats that were injected before consolidation and reconsolidation could take place, did not retain the fear response when they heard the tone again later. It seems that interference that is made before memories are consolidated affect the way they are remembered later.

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continually leading to strong connections between the two. Since the hippocampus can only support memories temporarily the remaining activation will be seen only in the neocortex which is able to support memory indefinitely. Squire and Alvarez took the temporally graded nature of patients with retrograde amnesia as support for the notion that once a connection has been established within the neocortex the hippocampus is no longer required, but this process is dynamic and extends for several years.

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in the functional magnetic resonance imaging have allowed them to improve their distinction between the hippocampus and the entorhinal cortex which they claim is more enduring in its activation from remote memory retrieval. They also criticize the use of memories during testing which cannot be confirmed as accurate. Finally, they state that the initial interview in the scanner acted as an encoding event as such differences between recent and remote memories would be obscured.

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debate and a detailed review of this field it had been concluded that reconsolidation was a real phenomenon. Tronson and Taylor compiled a lengthy summary of multiple reconsolidation studies, noting a number of studies were unable to show memory impairments due to blocked reconsolidation. However the need for standardized methods was underscored as in some learning tasks such as

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researchers have looked at the role growth hormones play in the consolidation of memories, particularly those of procedural and declarative memories. They found that although growth hormones support general brain systems and memory functioning, it is still unclear if growth hormones play a role in the formation and processing of particular memories during sleep periods.

78:. He noted the "curious fact... that the interval of a single night will greatly increase the strength of the memory," and presented the possibility that "... the power of recollection .. undergoes a process of ripening and maturing during the time which intervenes." The process of consolidation was later proposed based on clinical data illustrated in 1882 by 314:, which are suggested to underlie memory formation. LTP is also considered to be an important mechanism in terms of maintaining memories within brain regions, and therefore is thought to be involved in learning. There is compelling evidence that LTP is critical for Pavlovian fear conditioning in rats suggesting that it mediates 470:, representing the hippocampus-dependent stage. During this stage the hippocampus is 'teaching' the cortex more and more about the information and when the information is recalled it strengthens the cortico-cortical connection thus making the memory hippocampus-independent. Therefore, from one week and beyond the initial 85:, "progressive destruction advances progressively from the unstable to the stable". This idea was elaborated on by William H. Burnham a few years later in a paper on amnesia integrating findings from experimental psychology and neurology. Coining of the term "consolidation" is credited to the German researchers 236:(shown, for example, in goldfish), and as such it is considered the 'fast' type of consolidation. It is also referred to as 'initial consolidation'. As soon as six hours after training, memories become impervious to interferences that disrupt synaptic consolidation and the formation of long-term memory. 688:

regions; whereas the post-training awake group had no such improvements. It has been theorized that this may be related more-so to a process of synaptic consolidation rather than systems consolidation because of the short-term nature of the process involved. Researchers examining the effect of sleep

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related tasks. In one study testing finger-tapping, people were split into two groups and tested post-training with or without intervening sleep; results concluded that sleep post-training increases both speed and accuracy in this particular task, while increasing the activation of both cortical and

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however has been said to function separate from this system as it relies primarily on motor areas of the brain. The implicit nature of procedural knowledge allows it to exist absent from the conscious awareness that the information is there. Amnesic patients have shown retained ability to be trained

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memories no matter of their age. An important point they make while interpreting the results is that activation in the hippocampus was equally as strong regardless of the fact that the memories recalled were as old as 45 years prior to the date of the experiment. This is complicated by the fact that

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involved in the initial encoding of the memory. In this sense the MTL would act as a relay station for the various perceptual input that make up a memory and stores it as a whole event. After this has occurred the MTL directs information towards the neocortex to provide a permanent representation of

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have revolutionized the study of consolidation. Providing additional support is the study of functional brain activity in humans which has revealed that the activity of brain regions changes over time after a new memory is acquired. This change can occur as quickly as a couple hours after the memory

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More recent studies, however, have looked at the relationship between slow-wave sleep and memory consolidation, rather than REM sleep. One study found that low levels of acetylcholine found in the central nervous system, which are present during slow-wave sleep, aid in the consolidation of memories

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Haist, Gore, and Mao, sought to examine the temporal nature of consolidation within the hippocampus to test MTT against the standard view. They found that the hippocampus does not substantially contribute to the recollection of remote memories after a period of a few years. They claim that advances

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where it becomes permanently stored. In this view the hippocampus can perform the task of storing memories temporarily because the synapses are able to change quickly whereas the neocortical synapses change over time. Consolidation is thus the process whereby the hippocampus activates the neocortex

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for new consolidation, i.e., re-consolidation of the old memory. Nader, Schafe, and Le Doux (2000) demonstrated that the reconsolidation process may make memories more malleable than previously believed. Nader and his colleagues trained rats to be afraid of a tone by pairing the tone with a small

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need to be distinguished as relying on two different memory systems. When episodic information is encoded there are semantic aspects of the memory that are encoded as well and this is proposed as an explanation of the varying gradients of memory loss seen in amnesic patients. Amnesic patients with

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Some studies have supported this theory, while others have failed to demonstrate disruption of consolidated memory after retrieval. Negative results may be examples of conditions where memories are not susceptible to a permanent disruption, thus a determining factor of reconsolidation. After much

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One study that demonstrates this effect was conducted in 1984 by Smith and Rothkopf. In this experiment, subjects were sorted into three groups to test retention and learning. "Each group was taught the same 8 hour statistics class, but one group was taught the class in one day, the next over the

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have the opposite effect on the enhancement of memory consolidation. The BLA is thought to be actively involved in memory consolidation and is influenced strongly by stress hormones resulting in increased activation and as such increased memory retention. The BLA then projects to the hippocampus

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and addresses perceived shortcomings of the standard model with respect to the dependency of the hippocampus. MTT argues that the hippocampus is always involved in the retrieval and storage of episodic memories. It is thought that semantic memories, including basic information encoded during the

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In the decade between 2005 and 2015, at least five groups argued the notion that memory reconsolidation can be used to treat psychological problems. Three of these groups have proposed that the wide variety of different psychotherapies produce permanent change in clients to the extent that they

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suggesting that recently acquired memories of as long as a couple years could remain in the MTL prior to consolidation into other brain areas. Research into other patients with resections of the MTL have shown a positive relationship between the degree of memory impairment and the extent of MTL

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are a by-product of the reactivation of the brain areas and this can explain why dreams may be unrelated to the information being consolidated. The dream experience itself is not what enhances memory performance but rather it is the reactivation of the neural circuits that causes this. Other

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after they found that new information learned could disrupt information previously learnt if not enough time had passed to allow the old information to be consolidated. This led to the suggestion that new memories are fragile in nature but as time passes they become solidified.

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that lasts for at least 24 hours. Synaptic consolidation is achieved faster than systems consolidation (which is assumed to take weeks, months, or even to years to be accomplished). There is evidence to suggest that synaptic consolidation takes place within minutes to hours of

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in a more permanent form of storage. Systems consolidation is a slow dynamic process that can take anywhere from one to two decades to be fully formed in humans, unlike synaptic consolidation that only takes minutes to hours for new information to stabilize into memories.

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manage to activate this same neurobiological mechanism of reconsolidation in a way that leads to deconsolidation. One example of this is the Lefkoe Method, created in 1985 by Morty Lefkoe, president and founder of the Lefkoe Institute. Memory reconsolidation may be a

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Brunet, Alain; Orr, Scott P.; Tremblay, Jacques; Robertson, Kate; Nader, Karim; Pitman, Roger K. (2008). "Effect of post-retrieval propranolol on psychophysiologic responding during subsequent script-driven traumatic imagery in post-traumatic stress disorder".

272:. The result of the gene expression is the lasting alteration of synaptic proteins, as well as synaptic remodeling and growth. In a short time-frame immediately following learning, the molecular cascade, expression and process of both transcription factors and 789:(ECT). This seemed to indicate the involvement of a re-consolidation process for excited memories, and that the operation active in ECT was the disruption of that process; here, of the reconsolidation of retrieved fear memories by shock administration. 516:

the hippocampus is constantly involved in the encoding of new events and activation due to this fact is hard to separate using baseline measures. Because of this, activation of the hippocampus during retrieval of distant memories may simply be a

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is a category of processes that stabilize a memory trace after its initial acquisition. A memory trace is a change in the nervous system caused by memorizing something. Consolidation is distinguished into two specific processes. The first,

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activity following an enriched or novel waking experience, thus increasing neuronal plasticity and therefore playing an essential role in the consolidation of memories. This has come into question in recent years however and studies on

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have shown that animals and humans who are denied REM sleep do not show deficits in task learning. It has been proposed that since the brain is in a non-memory encoding state during sleep, consolidation would be unlikely to occur.

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by lengthening the reactivation phase. There have also been concerns about the use of reconsolidation research to justify psychotherapy treatments, and the generalizability of basic reconsolidation research into the therapy room

359:, and enhances relational memory consolidation. When interpreted in the context of synaptic consolidation, mechanisms of synaptic strengthening may depend on the spacing of memory reactivation to allow sufficient time for 354:

has been found to enhance memory consolidation, specifically for relational memory. Experimental results suggest that distributing learning over the course of 24 hours decreases the rate of forgetting compared to

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recall of facts, episodes, and lists, and its storage typically connected with the mediotemporal lobe and the hippocampal systems as it includes the encoding of both semantic and episodic information of events.

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and Alfons Pilzecker who rediscovered the concept that memory takes time to fixate or undergo "Konsolidierung" in their studies conducted between 1892 and 1900. The two proposed the perseveration-consolidation

621:. Studies appear to suggest that the amygdala effects the consolidation of memories through its influence with stress hormones and the projections to other brain areas implicated in memory consolidation. 394:. However, other results have shown that protein synthesis may not in fact be necessary for memory consolidation, as it has been found that the formation of memories can withstand vast amounts of 843:

In addition to fear memories, appetitive memories are also prone to reconsolidation episodes, which can likewise be disrupted; namely, after local administration of a protein activity inhibitor.

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that are involved in the information flow between relevant brain areas. A more complete understanding of these mechanics may possibly allow purposely enabling or strengthening this reactivation.

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Questions arose if reconsolidation was a unique process or merely another phase of consolidation. Both consolidation and reconsolidation can be disrupted by pharmacological agents (e.g. the

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Memory reconsolidation is the process of previously consolidated memories being recalled and actively consolidated. It is a distinct process that serves to maintain, strengthen and modify

390:, weaken memory, suggesting that protein synthesis is required for memory consolidation. Additionally, reports have suggested that the effects of protein synthesis inhibitors also inhibit 924:

are more difficult to run than typical consolidation experiments as disruption of a previously consolidated memory must be shown to be specific to the reactivation of the original

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Gais, Steffen; Hüllemann, Philipp; Hallschmid, Manfred; Born, Jan (2006). "Sleep-dependent surges in growth hormone do not contribute to sleep-dependent memory consolidation".

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was administered to the hippocampus, enhanced consolidation was seen during food-rewarded maze tasks. The opposite effect was also seen when the amygdala was inactivated using

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stable after retrieval is complete. It is believed that post-retrieval stabilization is different and distinct from consolidation, despite its overlap in function (e.g.

874:. Further studies have demonstrated an analogue of memory reconsolidation in spinal cord pain processing pathways, suggesting a general role for reconsolidation in the 147:

the patient began to suffer from memory impairments. Molaison lost the ability to encode and consolidate newly learned information leading researchers to conclude the

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Rasch, Björn H.; Born, Jan; Gais, Steffen (2006-05-01). "Combined Blockade of Cholinergic Receptors Shifts the Brain from Stimulus Encoding to Memory Consolidation".

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Gräff J; Joseph NF; Horn ME; Samiei A; Meng J; Seo J; Rei D; Bero AW; Phan TX; Wagner F; Holson E; Xu J; Sun J; Neve RL; Mach RH; Haggarty SJ; Tsai LH. (Jan 2014).

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The theory of reconsolidation has been debated for many years and is still controversial. Reconsolidation was first conceptualized in light of the discovery that

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Since those breakthrough studies were done, there have been several others to probe the theory of memory reconsolidation. Subjects in these studies, along with

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Nadel, L; Samsonovich, A; Ryan, L; Moscovitch, M (2000). "Multiple trace theory of human memory: Computational, neuroimaging, and neuropsychological results".

276:, are susceptible to disruptions. Disruptions caused by specific drugs, antibodies and gross physical trauma can block the effects of synaptic consolidation. 840:

These studies done by Nader and others seem to suggest that as memories are being remembered, they are fragile, as if experiencing them for the first time.

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Tse, D.; Langston, R. F.; Kakeyama, M.; Bethus, I.; Spooner, P. A.; Wood, E. R.; Witter, M. P.; Morris, R. G. M. (2007). "Schemas and Memory Consolidation".

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Okuda, K.; Højgaard, K.; Privitera, R.; Bayraktar, G.; Takeuchi, T. (2020). "Initial memory consolidation and the synaptic tagging and capture hypothesis".

902:. Under this possibility, traditional disruptions of reconsolidation might actually maintain the original memory trace but preventing the consolidation of 822:, but not by infusions made six hours afterwards. It was concluded that consolidated fear memory, when reactivated, enters a changeable state that requires 3576:

Centonze, Diego; Siracusano, Alberto; Calabresi, Paolo; Bernardi, Giorgio (October 2005). "Removing pathogenic memories: a neurobiology of psychotherapy".

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agent (leads to memory loss). These studies found it to be effective on retrieved memories when administered directly after the retrieval of a memory.

3213:"Activation of PKCzeta and PKMzeta in the nucleus accumbens core is necessary for the retrieval, consolidation and reconsolidation of the drug memory" 753:. Once memories undergo the process of consolidation and become part of long-term memory, they are thought of as stable. However, the retrieval of a 4796: 2235:

Gold, Paul E.; Van Buskirk, Roderick B. (February 1975). "Facilitation of time-dependent memory processes with posttrial epinephrine injections".

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The standard model of systems consolidation has been summarized by Squire and Alvarez (1995); it states that when novel information is originally

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Kindt, Merel; Soeter, Marieke; Vervliet, Bram (2009-02-15). "Beyond extinction: erasing human fear responses and preventing the return of fear".

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Recent studies have examined the relationship between REM sleep and procedural learning consolidation. In particular studies have been done on

134: 2369:"Involvement of amygdala pathways in the influence of post-training intra-amygdala norepinephrine and peripheral epinephrine on memory storage" 2094:

Haist, F.; Bowden Gore, J. B.; Mao, H. (2001). "Consolidation of human memory over decades revealed by functional magnetic resonance imaging".

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Nader, K.; Schafe, G. E.; LeDoux, J. E. (2000). "Fear memories require protein synthesis in the amygdala for reconsolidation after retrieval".

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Squire and Alvarez also proposed the idea that MTL structures play a role in the consolidation of memories within the neocortex by providing a

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Memory consolidation during sleep via reactivation of prior experiences and information is associated with sleep signatures of cortical "slow

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But for a more hesitant view of the role of memory reconsolidation in psychotherapy that criticizes some of the claims of Ecker et al., see:

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is not just due to task impairment caused by the procedure, which can be demonstrated by testing control groups in absence of the original

31:, occurs on a small scale in the synaptic connections and neural circuits within the first few hours after learning. The second process is 4379: 5574: 5245: 1399:

Milner, B.; Corkin, S.; Teuber, H. -L. (1968). "Further analysis of the hippocampal amnesic syndrome: 14-year follow-up study of H.M".

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McIntyre, C. K.; Power, A. N. E.; Roozendaal, B.; McGaugh, J. L. (2006). "Role of the Basolateral Amygdala in Memory Consolidation".

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Walker, M.P.; Stickgold, R.; Alsop, D.; Gaab, N.; Schlaug, G. (2005). "Sleep-dependent motor memory plasticity in the human brain".

737:, suggest that a waking experience prior to sleep can have an enduring effect in the brain, due to an increase of neuroplasticity. 3146:

Schiller, Daniela; Monfils, Marie-H.; Raio, Candace M.; Johnson, David C.; LeDoux, Joseph E.; Phelps, Elizabeth A. (2009-12-09).

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modifications may also prevent reconsolidation in some cases. The removal of these epigenetic modifications with inhibitors of

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Bramham, C. R.; Messaoudi, E. (2005). "BDNF function in adult synaptic plasticity: The synaptic consolidation hypothesis".

4210: 4870: 3696:"Memory reconsolidation, emotional arousal and the process of change in psychotherapy: new insights from brain science" 1052: 1580:"Food for thought: The role of dietary flavonoids in enhancing human memory, learning and neuro-cognitive performance" 3822: 1723:

Smith, Steven M.; Rothkopf, Ernst Z. (1984). "Contextual Enrichment and Distribution of Practice in the Classroom".

773:). Memory modification needs to be demonstrated in the retrieval in order for this independent process to be valid. 4255: 2192:

McGaugh, J. L.; Roozendaal, B. (2002). "Role of adrenal stress hormones in forming lasting memories in the brain".

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will block the retention of memory effects seen previously. This is supported by the fact that beta-adrenoreceptor

398:, suggesting that this criterion of protein synthesis as necessary for memory consolidation is not unconditional. 3695: 2957:

Solyom, L.; Kenny, F.; Ledwidge, B. (1969). "Psychotherapy: Evaluation of a New Treatment Paradigm for Phobias".

1932:

Squire, L. R.; Alvarez, P. (1995). "Retrograde amnesia and memory consolidation: A neurobiological perspective".

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after pre-exposure to an enriched environment. Results from studies testing the effects of zif268 on mice brains

2422:"The basolateral amygdala is a cofactor in memory enhancement produced by intrahippocampal glutamate injections" 5238: 4743: 3703: 834: 792:

Further studies investigated the concept, using ECT to test for reconsolidation; ECT was already known as an

5345: 3529:"A requirement for the immediate early gene zif268 in reconsolidation of recognition memory after retrieval" 2316:"Modulating effects of posttraining epinephrine on memory: involvement of the amygdala noradrenergic system" 4791: 4692: 4579: 1620: 833:

Brunet and colleagues (2008) studied patients that had been diagnosed with Post Traumatic Stress Disorder (

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Nadel, L.; Moscovitch, M. (1997). "Memory consolidation, retrograde amnesia and the hippocampal complex".

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resulting in a strengthened memory. This relationship was studied by Packard and Chen who found that when

237: 28: 5340: 395: 383: 3413:"Directly reactivated, but not indirectly reactivated, memories undergo reconsolidation in the amygdala" 2884:"Endogenous memory reactivation during sleep in humans is clocked by slow oscillation-spindle complexes" 4917: 4842: 4675: 3000:

Nader, Karim; Schafe, Glenn E.; LeDoux, Joseph E. (2000). "The labile nature of consolidation theory".

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Wilhelm, Ines; Diekelmann, Susanne; Molzow, Ina; Ayoub, Amr; Mölle, Matthias; Born, Jan (2011-02-02).

5231: 3966:"Hippocampal replay in the awake state: A potential substrate for memory consolidation and retrieval" 1040: 4029: 2857: 2478: 5407: 5268: 4955: 4900: 4875: 4705: 4682: 4632: 4537: 4201: 1062: 79: 5528: 4642: 4408: 3922: 3368:

Patihis, L. (2015). "Let's be skeptical about reconsolidation and emotional arousal in therapy".

949: 903: 892: 299: 176: 3321:"Epigenetic priming of memory updating during reconsolidation to attenuate remote fear memories" 2279: 2271: 1067: 5498: 5300: 5049: 5009: 4910: 4879: 4517: 4305: 4196: 4024: 3810:

Unlocking the Emotional Brain: Eliminating Symptoms at Their Roots Using Memory Reconsolidation

3578: 2473: 965: 875: 391: 303: 285: 4064:"Dreaming of a Learning Task is Associated with Enhanced Sleep-Dependent Memory Consolidation" 3397: 2743:"Dreaming of a Learning Task is Associated with Enhanced Sleep-Dependent Memory Consolidation" 2368: 2315: 5548: 5513: 5293: 5278: 5064: 4779: 4665: 4637: 4622: 4617: 4455: 1036: 988: 436: 291: 273: 265: 43:

over a period of weeks to years. Recently, a third process has become the focus of research,

86: 4948: 4932: 4811: 4569: 4522: 4512: 4248: 4075: 4016: 3760: 3483: 3424: 3224: 3211:

Crespo, J.A.; Stöckl, P.; Ueberall, F.; Marcel, J.; Saria, A.; Zernig, G. (February 2012).

3159: 2895: 2754: 2155: 1575: 1311: 1205: 1123: 984: 973: 722: 601: 589: 572: 351: 148: 8: 5543: 5079: 4979: 4670: 4554: 4502: 4470: 4450: 3472:"Independent cellular processes for hippocampal memory consolidation and reconsolidation" 2882:

Schreiner, Thomas; Petzka, Marit; Staudigl, Tobias; Staresina, Bernhard P. (2021-05-25).

1978:

Frankland, P. W.; Bontempi, B. (2005). "The organization of recent and remote memories".

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Tronson, N. C.; Taylor, J. R. (2007). "Molecular mechanisms of memory reconsolidation".

1315: 1209: 223:, when discussed in the context of synaptic consolidation, is conventionally said to be 219:

is one form of memory consolidation seen across all species and long-term memory tasks.

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McGaugh, J. L. (2002). "Memory consolidation and the amygdala: A systems perspective".

4096: 4063: 4050: 3990: 3965: 3869: 3781: 3746: 3603: 3558: 3509: 3447: 3412: 3345: 3320: 3296: 3271: 3247: 3212: 3188: 3147: 3128: 3033: 2924: 2883: 2834: 2799: 2775: 2742: 2718: 2667: 2613: 2499: 2487: 2443: 2399: 2346: 2217: 2119: 2069: 2003: 1957: 1874: 1849: 1797: 1661: 1621:"Long-term potentiation in the amygdala: A mechanism for emotional learning and memory" 1557: 1495: 1229: 1173: 1110:

Dudai, Y. (2004). "The Neurobiology of Consolidations, Or, How Stable is the Engram?".

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in an effort to identify brain substrates critical for slow consolidation. Meanwhile,

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Nadel and Moscovitch argued that when studying the structures and systems involved in

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regions. Later the hippocampus' representations of this information become active in

432: 415: 375: 360: 330: 311: 253: 197: 188: 82: 4226: 4142: 4054: 3607: 3562: 3513: 2671: 2350: 2221: 2123: 1801: 1665: 1177: 1161: 5533: 5472: 5387: 5151: 5104: 5074: 5029: 4885: 4816: 4769: 4574: 4549: 4435: 4395: 4283: 4206: 4179: 4159: 4122: 4091: 4083: 4034: 3985: 3977: 3853: 3776: 3768: 3751: 3742: 3738: 3712: 3665: 3587: 3540: 3491: 3442: 3432: 3377: 3340: 3332: 3291: 3283: 3242: 3232: 3183: 3167: 3132: 3104: 3061: 3037: 3009: 2966: 2919: 2903: 2829: 2816: 2811: 2770: 2762: 2722: 2694: 2647: 2643: 2595: 2543: 2535: 2503: 2483: 2433: 2403: 2383: 2330: 2283: 2244: 2201: 2163: 2103: 2073: 2053: 2007: 1987: 1961: 1941: 1869: 1861: 1771: 1732: 1692: 1643: 1635: 1591: 1561: 1541: 1479: 1442: 1408: 1371: 1363: 1319: 1263: 1233: 1213: 1157: 1119: 766: 750: 630: 512: 295: 252:

The standard model of synaptic consolidation suggests that alterations of synaptic

220: 4062:

Wamsley, E. J.; Tucker, M.; Payne, J. D.; Benavides, J. A.; Stickgold, R. (2010).

2741:

Wamsley, E. J.; Tucker, M.; Payne, J. D.; Benavides, J. A.; Stickgold, R. (2010).

2629: 2627: 2617: 932:

of reactivation occurs in a limited time frame, which can be assessed by delaying

98: 5569: 5477: 5422: 5392: 5380: 5335: 5320: 5089: 5069: 5044: 5034: 4989: 4984: 4738: 4710: 4445: 4428: 4423: 4418: 4413: 4288: 4241: 3808: 3747:"Preventing the return of fear in humans using reconsolidation update mechanisms" 3272:"A spinal analogue of memory reconsolidation enables the erasure of hyperalgesia" 3237: 3148:"Preventing the return of fear in humans using reconsolidation update mechanisms" 2600: 2583: 1901: 1819: 1447: 1430: 1323: 863: 651: 546: 542: 507: 503: 487: 483: 452: 448: 444: 269: 229: 183:

possibly responsible for fast consolidation. In recent decades, advancements in

5156: 5120: 5014: 4612: 4559: 4385: 4355: 4335: 4322: 3336: 2970: 2944: 2907: 2698: 2624: 356: 346: 184: 111: 4087: 3912: 3716: 3591: 3381: 2766: 1865: 1760:"Effects of spacing and embellishment on memory for the main points of a text" 1736: 1596: 1579: 5563: 5432: 5310: 5305: 5135: 5125: 5099: 5094: 5054: 5039: 5004: 4927: 4774: 4602: 4465: 4440: 4403: 4360: 4350: 4345: 4330: 3857: 3832: 3691: 3179: 3116: 3073: 3021: 2978: 2915: 2858:"Press (re)play to remember: How the brain strengthens memories during sleep" 2825: 2706: 2655: 1785: 1744: 933: 929: 726: 707: 323: 192: 137: 4038: 3496: 3471: 3437: 2167: 102:

Lateral view of the hippocampus which is located in the medial temporal lobe

5375: 5166: 5130: 5084: 4994: 4837: 4652: 4607: 4594: 4584: 4544: 4264: 4218: 4171: 4134: 4105: 4046: 3999: 3865: 3790: 3724: 3599: 3554: 3505: 3456: 3389: 3354: 3305: 3256: 3197: 3124: 3081: 3029: 2933: 2843: 2784: 2714: 2663: 2609: 2557: 2524:"Brain Gene Expression During REM Sleep Depends on Prior Waking Experience" 2495: 2213: 2115: 1999: 1883: 1706: 1657: 1605: 1553: 1491: 1456: 1385: 1331: 1277: 1225: 1169: 1131: 925: 899: 799:

Later research, wherein fear memories had been established in rats through

754: 244:, is thought to be the cellular process underlying synaptic consolidation. 68: 67:

Memory consolidation was first referred to in the writings of the renowned

2986: 2395: 2342: 2256: 2175: 2065: 1953: 1793: 948:. Finally, it is important to rule out alternative explanations, such as 204:

dimension to the reorganization of the memory as it is represented in the

5352: 5325: 5288: 5024: 4895: 4849: 3649:"Transformative emotional sequence: towards a common principle of change" 3623:"Unlocking the emotional brain: finding the neural key to transformation" 1648: 1367: 1007: 910: 685: 680: 655: 647: 605: 463: 440: 411: 410:

consolidation. It is a reorganization process in which memories from the

171:

These studies were accompanied by the creation of animal models of human

123: 75: 40: 5523: 3772: 3171: 2270:

Gold, P. E.; McIntyre, C.; McNay, E.; Stefani, M.; Korol, D. L. (2001).

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Cognitive Psychology: Connecting Mind, Research and Everyday Experience

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till six hours after reactivation. It is also useful to show that the

921: 819: 734: 593: 517: 475: 419: 91: 54: 3887: 3210: 2800:"Sleep Selectively Enhances Memory Expected to Be of Future Relevance" 2287: 1483: 1268: 1251: 5508: 5493: 5330: 5273: 5260: 4295: 3936: 3814: 3669: 3013: 1252:"Retrieval and reconsolidation: toward a neurobiology of remembering" 1217: 859: 800: 730: 635: 618: 614: 567: 553: 180: 152: 3981: 3287: 3108: 1991: 1545: 983:

is required for consolidation (but not reconsolidation) whereas the

891:, certain forms of memory reactivation could actually represent new 5518: 5427: 5223: 4784: 3575: 952: 945: 906: 895: 811: 746: 697: 659: 643: 609: 585: 471: 467: 387: 379: 334: 315: 233: 165: 144: 115: 72: 48: 2107: 1718: 1716: 994:

is required for reconsolidation but not consolidation. A similar

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10.1002/1098-1063(2000)10:4<352::AID-HIPO2>3.0.CO;2-D

3917: 871: 867: 855: 793: 782: 490: 216: 172: 160: 127: 119: 2881: 642:

has been thought of to be an important concept in the overnight

5454: 4233: 3318: 1850:"Protein synthesis inhibition and memory: Formation vs amnesia" 1713: 1681:"Distributed learning enhances relational memory consolidation" 1469: 1015: 999: 991: 941: 807: 762: 758: 718: 579: 538: 407: 319: 240:, the long-lasting form of one of the best understood forms of 224: 4186: 4149: 502:

Multiple trace theory (MTT) builds on the distinction between

5401: 2939: 1302:

McGaugh, J. L. (2000). "Memory--a Century of Consolidation".

851: 847: 690: 639: 456: 205: 2797: 806:, found that a consolidated fear memory can be brought to a 4061: 2740: 2633: 1352:"Loss of Recent Memory After Bilateral Hippocampal Lesions" 1003: 980: 976: 201: 179:

studies of selected brain areas began to shed light on the

35:, occurring on a much larger scale in the brain, rendering 3741:; Monfils, Marie-H.; Raio, Candace M.; Johnson, David C.; 3737: 3145: 3050: 302:

sensitivity, lasting minutes to even days. The process of

4006: 3913:"Eliminate Core Beliefs - Morty Lefkoe on the Today Show" 2276:

Memory consolidation: Essays in honor of James L. McGaugh

558:

Learning can be distinguished by two forms of knowledge:

3411:

Debiec, J.; Doyere, V.; Nader, K.; LeDoux, J.E. (2006).

3269: 3158:(7277). Springer Science and Business Media LLC: 49–53. 2463: 2269: 1899: 928:. Furthermore, it is important to demonstrate that the 290:

LTP can be thought of as the prolonged strengthening of

155:

involved in this process. Molaison also showed signs of

47:, in which previously consolidated memories can be made 3410: 3103:(3). Springer Science and Business Media LLC: 256–258. 3008:(3). Springer Science and Business Media LLC: 216–219. 761:

phase that then requires an active process to make the

130:

and their effect on memory. After Molaison underwent a

3685: 1770:(2). Springer Science and Business Media LLC: 97–102. 725:(IEG) thought to be involved in neuroplasticity by an 110:

started to emerge in the 1960s and 1970s. The case of

1900:

Roediger, H.L.; Dudai, Y.; Fitzpatrick, S.M. (2007).

917:

enabled the erasure of remote memories after recall.

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experience, the memory is slowly transferred to the

159:

spanning a period of about three years prior to the

3807:Ecker, Bruce; Ticic, Robin; Hulley, Laurel (2012). 3094: 2956: 2093: 2039: 2037: 1424: 1422: 1356:

Journal of Neurology, Neurosurgery & Psychiatry

1029: 363:to occur, and thereby strengthen long-term memory. 3845:Journal of the American Psychoanalytic Association 3470:Lee, J. L.; Everitt, B. J.; Thomas, K. L. (2004). 2272:"Neurochemical referees of dueling memory systems" 2035: 2033: 2031: 2029: 2027: 2025: 2023: 2021: 2019: 2017: 1398: 979:. However, recent amygdala research suggests that 959: 554:Declarative vs. procedural knowledge consolidation 3964:Carr, M. F.; Jadhav, S. P.; Frank, L. M. (2011). 3526: 2999: 2420:Packard, Mark G; Chen, Scott A (September 1999). 2367:Liang, KC; McGaugh, JL; Yao, HY (February 1990). 2313: 2191: 2187: 2185: 1977: 1813: 1811: 1195: 5561: 3806: 3469: 2314:Liang, KC; Juler, RG; McGaugh, JL (March 1986). 1463: 1431:"The consolidation and transformation of memory" 1419: 1345: 1343: 1341: 1147: 698:Cortical slow oscillation- and spindle-complexes 378:plays an important role in the formation of new 337:synaptic transmission that would result in LTP. 51:again through reactivation of the memory trace. 3963: 2366: 2234: 2043: 2014: 1297: 1295: 1293: 1291: 1289: 1287: 532: 520:of the subject encoding the study as an event. 3642: 3640: 2228: 2182: 2146:Squire, L. R. (1986). "Mechanisms of memory". 1927: 1925: 1923: 1808: 1428: 1349: 1143: 1141: 672:and therefore assist in the learning process. 329:appear to block the induction of both LTP and 5239: 4249: 3888:"How the Lefkoe Belief Process works, Part 1" 2684: 1931: 1757: 1722: 1678: 1531: 1338: 646:in humans by establishing information in the 126:and sparked massive interest in the study of 58:The line processes to make information memory 3802: 3800: 1973: 1971: 1284: 580:Emotional and stressful memory consolidation 406:Systems consolidation is the second form of 4380:The Magical Number Seven, Plus or Minus Two 3681: 3679: 3637: 3204: 1920: 1895: 1893: 1758:Reder, Lynne M.; Anderson, John R. (1982). 1138: 294:, and is known to produce increases in the 114:, formerly known as patient H.M., became a 5246: 5232: 4256: 4242: 4152:Annals of the New York Academy of Sciences 3527:Bozon, B.; Davis, S.; Laroche, S. 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Declarative information includes the 5562: 2581: 2564: 2454: 2357: 2145: 2130: 1672: 1245: 1243: 1124:10.1146/annurev.psych.55.090902.142050 785:could often be eliminated by means of 624: 118:in studies of memory as it relates to 5227: 4237: 3620: 2304: 2263: 2080: 1618: 1612: 1392: 1109: 940:measure used to assess disruption of 333:and that fear conditioning increases 5253: 3842:(April 2015). "Commentary on Tuch". 3657:Journal of Psychotherapy Integration 3312: 1847: 1838: 1584:Proceedings of the Nutrition Society 1350:Scoville, W. B.; Milner, B. (1957). 1249: 1184: 1084: 435:and registered, memory of these new 370: 27:, which is thought to correspond to 1854:Neurobiology of Learning and Memory 1240: 1006:for consolidation was found in the 729:of the transcription factor during 658:. REM sleep elicits an increase in 426: 264:. These molecular cascades trigger 262:intracellular transduction cascades 164:removal which points to a temporal 13: 4164:10.1111/j.1749-6632.2003.tb07088.x 3957: 2488:10.1016/j.neuroscience.2005.04.007 1824:(4th ed.). Cengage Learning. 1731:(3). Informa UK Limited: 341–358. 740: 14: 5586: 5575:Unsolved problems in neuroscience 4661:Deese–Roediger–McDermott paradigm 2687:Journal of Cognitive Neuroscience 909:. Recent work has suggested that 898:rather than activation of an old 466:for up to one week after initial 414:region, where memories are first 340: 247: 29:late-phase long-term potentiation 5206: 5194: 4263: 3925:from the original on 2021-12-15. 3066:10.1016/j.jpsychires.2007.05.006 2938: 1679:Litman, L.; Davachi, L. (2008). 1472:European Journal of Neuroscience 1030:Reconsolidation in psychotherapy 260:are achieved through activating 3929: 3905: 3880: 3731: 3621:Ecker, Bruce (September 2008). 3614: 3569: 3520: 3463: 3404: 3361: 3139: 3088: 3054:Journal of Psychiatric Research 3044: 2993: 2950: 2875: 2850: 2791: 2678: 2584:"Memory Consolidation in Sleep" 2194:Current Opinion in Neurobiology 2046:Current Opinion in Neurobiology 1934:Current Opinion in Neurobiology 1751: 1162:10.1016/j.pneurobio.2005.06.003 960:Distinctions from consolidation 881: 787:electroconvulsive shock therapy 749:that are already stored in the 4871:Atkinson–Shiffrin memory model 4744:Memory and social interactions 2817:10.1523/JNEUROSCI.3575-10.2011 2648:10.1016/j.psyneuen.2006.02.009 1053:Atkinson–Shiffrin memory model 1: 4127:10.1016/S0166-2236(02)02211-7 3704:Behavioral and Brain Sciences 3686:Lane, Richard D.; Ryan, Lee; 3546:10.1016/s0896-6273(03)00674-3 3370:Behavioral and Brain Sciences 2249:10.1016/S0091-6773(75)91784-8 2206:10.1016/S0959-4388(02)00306-9 2058:10.1016/S0959-4388(97)80010-4 1640:10.1016/S0166-2236(99)01465-4 1078: 810:state, by means of immediate 523: 455:(unconscious) recall like in 439:becomes retained in both the 5408:Perception as interpretation 4580:Retrieval-induced forgetting 3238:10.1371/journal.pone.0030502 2601:10.1016/j.neuron.2004.08.034 2388:10.1016/0006-8993(90)90400-6 2335:10.1016/0006-8993(86)91049-8 1946:10.1016/0959-4388(95)80023-9 1448:10.1016/j.neuron.2015.09.004 1413:10.1016/0028-3932(68)90021-3 1324:10.1126/science.287.5451.248 776: 533:Semantic vs. episodic memory 396:protein synthesis inhibition 384:protein synthesis inhibitors 39:memories independent of the 7: 3647:Welling, Hans (June 2012). 3060:(6). Elsevier BV: 503–506. 3002:Nature Reviews Neuroscience 2642:(6). Elsevier BV: 786–791. 1980:Nature Reviews Neuroscience 1906:. Oxford University Press. 1903:Science of Memory: Concepts 1534:Nature Reviews Neuroscience 1112:Annual Review of Psychology 1073:Sharp wave–ripple complexes 1046: 1041:many forms of psychotherapy 769:) and its mechanisms (e.g. 200:suggesting that there is a 10: 5591: 4918:Levels of Processing model 4843:World Memory Championships 4676:Lost in the mall technique 4523:dissociative (psychogenic) 3337:10.1016/j.cell.2013.12.020 2971:10.1177/070674376901400102 2908:10.1038/s41467-021-23520-2 2699:10.1162/jocn.2006.18.5.793 628: 462:Memory is retained in the 382:. Studies have shown that 344: 322:in mammals. Specifically, 283: 62: 5486: 5453: 5361: 5259: 5189: 5144: 5113: 4972: 4965: 4858: 4830: 4762: 4719: 4691: 4651: 4593: 4488: 4394: 4369: 4321: 4314: 4271: 4088:10.1016/j.cub.2010.03.027 3717:10.1017/S0140525X14000041 3382:10.1017/S0140525X14000272 2767:10.1016/j.cub.2010.03.027 1866:10.1016/j.nlm.2007.10.006 1737:10.1207/s1532690xci0103_4 1725:Cognition and Instruction 1597:10.1017/S0029665108007088 459:and 'offline' processes. 4956:The Seven Sins of Memory 4901:Intermediate-term memory 4706:Indirect tests of memory 4683:Recovered-memory therapy 4633:Misattribution of memory 3858:10.1177/0003065115579720 2636:Psychoneuroendocrinology 1818:Goldstein, E.B. (2014). 1150:Progress in Neurobiology 1002:for reconsolidation and 268:that lead to changes in 5529:Relational frame theory 5504:Higher nervous activity 4643:Source-monitoring error 4115:Trends in Neurosciences 4039:10.1126/science.1135935 3627:Psychotherapy Networker 3497:10.1126/science.1095760 3438:10.1073/pnas.0507168103 2804:Journal of Neuroscience 2168:10.1126/science.3086978 1628:Trends in Neurosciences 972:) and both require the 151:(MTL) was an important 122:and the removal of the 5499:Experiential avoidance 5050:George Armitage Miller 5010:Patricia Goldman-Rakic 3579:Molecular Neurobiology 2582:Vertes, R. P. (2004). 1764:Memory & Cognition 876:central nervous system 714:Zif268 & REM sleep 451:(conscious) recall or 286:Long-term potentiation 280:Long-term potentiation 217:Synaptic consolidation 212:Synaptic consolidation 106:Systematic studies of 103: 59: 25:synaptic consolidation 5514:Ironic process theory 5279:Cognitive flexibility 5213:Philosophy portal 5201:Psychology portal 5065:Henry L. Roediger III 4666:False memory syndrome 4638:Misinformation effect 4618:Imagination inflation 3840:Alberini, Cristina M. 2888:Nature Communications 2528:Learning & Memory 598:beta-andrenoreceptors 498:Multiple trace theory 402:Systems consolidation 310:and in the growth of 292:synaptic transmission 274:immediate early genes 266:transcription factors 101: 57: 37:hippocampus-dependent 33:systems consolidation 4570:Motivated forgetting 3941:The Lefkoe Institute 3937:"About Morty Lefkoe" 3892:The Lefkoe Institute 3692:Greenberg, Leslie S. 2522:Ribeiro, S. (1999). 1848:Gold, P. E. (2008). 1368:10.1136/jnnp.20.1.11 989:immediate early gene 985:transcription factor 974:transcription factor 723:Immediate early gene 573:Procedural knowledge 352:Distributed learning 177:neuropharmacological 149:medial temporal lobe 20:Memory consolidation 5544:Thought suppression 5080:Arthur P. Shimamura 4980:Richard C. Atkinson 4797:Effects of exercise 4671:Memory implantation 4555:Interference theory 4471:Selective retention 4451:Meaningful learning 4080:2010CBio...20..850W 4021:2007Sci...316...76T 3970:Nature Neuroscience 3773:10.1038/nature08637 3765:2010Natur.463...49S 3592:10.1385/MN:32:2:123 3488:2004Sci...304..839L 3429:2006PNAS..103.3428D 3229:2012PLoSO...730502C 3172:10.1038/nature08637 3164:2010Natur.463...49S 3097:Nature Neuroscience 2900:2021NatCo..12.3112S 2759:2010CBio...20..850W 2160:1986Sci...232.1612S 2154:(4758): 1612–1619. 2096:Nature Neuroscience 1316:2000Sci...287..248M 1210:2000Natur.406..722N 1020:double dissociation 1014:. However, not all 996:double dissociation 915:histone deacetylase 625:Sleep consolidation 588:, specifically the 418:, are moved to the 386:administered after 308:synaptic plasticity 242:synaptic plasticity 108:anterograde amnesia 5177:Andriy Slyusarchuk 5000:Hermann Ebbinghaus 4906:Involuntary memory 4807:Memory improvement 4792:Effects of alcohol 4754:Transactive memory 4732:Politics of memory 4701:Exceptional memory 2540:10.1101/lm.6.5.500 2439:10.3758/BF03332131 2237:Behavioral Biology 1777:10.3758/bf03209210 1698:10.1101/lm.1132008 1619:Maren, S. (1999). 1024:object recognition 757:can cause another 636:Rapid eye movement 590:basolateral region 486:area for multiple 258:membrane potential 157:retrograde amnesia 145:epileptic symptoms 104: 60: 5557: 5556: 5316:Critical thinking 5284:Cognitive liberty 5221: 5220: 5185: 5184: 5172:Cosmos Rossellius 5020:Marcia K. Johnson 4891:Exosomatic memory 4876:Context-dependent 4866:Absent-mindedness 4749:Memory conformity 4727:Collective memory 4628:Memory conformity 4565:Memory inhibition 4484: 4483: 4476:Tip of the tongue 3943:. 18 January 2008 3743:LeDoux, Joseph E. 3739:Schiller, Daniela 3482:(5672): 839–843. 2862:medicalxpress.com 2297:978-1-55798-783-9 2288:10.1037/10413-012 2102:(11): 1139–1145. 1913:978-0-19-972751-3 1831:978-1-305-17699-7 1576:Spencer, J. P. E. 1484:10.1111/ejn.14902 1310:(5451): 248–251. 1269:10.1101/lm.7.2.73 1250:Sara, SJ (2000). 1204:(6797): 722–726. 1058:Coherence therapy 1012:fear conditioning 966:protein synthesis 889:fear conditioning 827:protein synthesis 816:protein synthesis 814:infusions of the 804:fear conditioning 771:protein synthesis 665:sleep deprivation 376:Protein synthesis 371:Protein synthesis 361:protein synthesis 331:fear conditioning 312:synaptic strength 254:protein synthesis 189:molecular biology 83:Law of Regression 5582: 5254:Mental processes 5248: 5241: 5234: 5225: 5224: 5211: 5210: 5209: 5199: 5198: 5197: 5152:Jonathan Hancock 5105:Robert Stickgold 5075:Richard Shiffrin 5030:Elizabeth Loftus 4970: 4969: 4886:Childhood memory 4693:Research methods 4575:Repressed memory 4550:Forgetting curve 4538:transient global 4409:Autobiographical 4319: 4318: 4258: 4251: 4244: 4235: 4234: 4230: 4204: 4183: 4146: 4109: 4099: 4058: 4032: 4003: 3993: 3952: 3951: 3949: 3948: 3933: 3927: 3926: 3909: 3903: 3902: 3900: 3899: 3884: 3878: 3877: 3836: 3804: 3795: 3794: 3784: 3735: 3729: 3728: 3700: 3683: 3674: 3673: 3670:10.1037/a0027786 3653: 3644: 3635: 3634: 3618: 3612: 3611: 3573: 3567: 3566: 3548: 3524: 3518: 3517: 3499: 3467: 3461: 3460: 3450: 3440: 3423:(9): 3428–3433. 3408: 3402: 3401: 3365: 3359: 3358: 3348: 3331:(1–2): 261–276. 3316: 3310: 3309: 3299: 3282:(8): 1043–1045. 3267: 3261: 3260: 3250: 3240: 3208: 3202: 3201: 3191: 3143: 3137: 3136: 3092: 3086: 3085: 3048: 3042: 3041: 3014:10.1038/35044580 2997: 2991: 2990: 2954: 2948: 2943:Available under 2942: 2937: 2927: 2879: 2873: 2872: 2870: 2868: 2854: 2848: 2847: 2837: 2819: 2810:(5): 1563–1569. 2795: 2789: 2788: 2778: 2738: 2727: 2726: 2682: 2676: 2675: 2631: 2622: 2621: 2603: 2579: 2562: 2561: 2551: 2519: 2508: 2507: 2481: 2461: 2452: 2451: 2441: 2417: 2408: 2407: 2373: 2364: 2355: 2354: 2320: 2311: 2302: 2301: 2267: 2261: 2260: 2232: 2226: 2225: 2189: 2180: 2179: 2143: 2128: 2127: 2091: 2078: 2077: 2041: 2012: 2011: 1975: 1966: 1965: 1929: 1918: 1917: 1897: 1888: 1887: 1877: 1845: 1836: 1835: 1815: 1806: 1805: 1779: 1755: 1749: 1748: 1720: 1711: 1710: 1700: 1676: 1670: 1669: 1651: 1625: 1616: 1610: 1609: 1599: 1572: 1566: 1565: 1529: 1504: 1503: 1478:(8): 6826–6849. 1467: 1461: 1460: 1450: 1426: 1417: 1416: 1401:Neuropsychologia 1396: 1390: 1389: 1379: 1347: 1336: 1335: 1299: 1282: 1281: 1271: 1247: 1238: 1237: 1218:10.1038/35021052 1193: 1182: 1181: 1145: 1136: 1135: 1107: 1018:tasks show this 920:Reconsolidation 850:, have included 751:long-term memory 631:Sleep and memory 513:autobiographical 296:neurotransmitter 221:Long-term memory 124:hippocampal zone 5590: 5589: 5585: 5584: 5583: 5581: 5580: 5579: 5560: 5559: 5558: 5553: 5482: 5449: 5357: 5336:Problem solving 5321:Decision-making 5255: 5252: 5222: 5217: 5207: 5205: 5195: 5193: 5181: 5162:Dominic O'Brien 5140: 5109: 5090:Susumu Tonegawa 5070:Daniel Schacter 5045:Eleanor Maguire 5035:Geoffrey Loftus 4990:Stephen J. Ceci 4985:Robert A. Bjork 4961: 4880:state-dependent 4854: 4826: 4758: 4739:Cultural memory 4715: 4711:Memory disorder 4687: 4647: 4589: 4480: 4390: 4365: 4310: 4267: 4262: 4068:Current Biology 4030:10.1.1.385.8987 4015:(5821): 76–82. 3982:10.1038/nn.2732 3960: 3958:Further reading 3955: 3946: 3944: 3935: 3934: 3930: 3921:. 7 July 2009. 3911: 3910: 3906: 3897: 3895: 3886: 3885: 3881: 3825: 3805: 3798: 3759:(7277): 49–53. 3736: 3732: 3698: 3684: 3677: 3651: 3645: 3638: 3619: 3615: 3574: 3570: 3525: 3521: 3468: 3464: 3409: 3405: 3366: 3362: 3317: 3313: 3288:10.1038/nn.3758 3268: 3264: 3209: 3205: 3144: 3140: 3109:10.1038/nn.2271 3093: 3089: 3049: 3045: 2998: 2994: 2955: 2951: 2880: 2876: 2866: 2864: 2856: 2855: 2851: 2796: 2792: 2747:Current Biology 2739: 2730: 2683: 2679: 2632: 2625: 2580: 2565: 2520: 2511: 2479:10.1.1.471.2164 2462: 2455: 2418: 2411: 2371: 2365: 2358: 2318: 2312: 2305: 2298: 2268: 2264: 2233: 2229: 2190: 2183: 2144: 2131: 2092: 2081: 2042: 2015: 1992:10.1038/nrn1607 1976: 1969: 1930: 1921: 1914: 1898: 1891: 1846: 1839: 1832: 1816: 1809: 1756: 1752: 1721: 1714: 1677: 1673: 1634:(12): 561–567. 1623: 1617: 1613: 1573: 1569: 1546:10.1038/nrn2090 1530: 1507: 1468: 1464: 1427: 1420: 1397: 1393: 1348: 1339: 1300: 1285: 1248: 1241: 1194: 1185: 1146: 1139: 1108: 1085: 1081: 1049: 1032: 962: 884: 779: 743: 741:Reconsolidation 716: 700: 654:regions of the 633: 627: 582: 556: 547:episodic memory 543:semantic memory 541:consolidation, 535: 526: 508:episodic memory 504:semantic memory 500: 429: 404: 373: 357:massed learning 349: 343: 298:production and 288: 282: 270:gene expression 256:and changes in 250: 230:memory encoding 214: 65: 45:reconsolidation 17: 12: 11: 5: 5588: 5578: 5577: 5572: 5555: 5554: 5552: 5551: 5546: 5541: 5536: 5531: 5526: 5524:Mental fatigue 5521: 5516: 5511: 5506: 5501: 5496: 5490: 5488: 5484: 5483: 5481: 5480: 5475: 5470: 5465: 5459: 5457: 5451: 5450: 5448: 5447: 5442: 5441: 5440: 5435: 5430: 5420: 5415: 5410: 5405: 5395: 5390: 5385: 5384: 5383: 5373: 5367: 5365: 5359: 5358: 5356: 5355: 5350: 5349: 5348: 5343: 5333: 5328: 5323: 5318: 5313: 5308: 5303: 5298: 5297: 5296: 5286: 5281: 5276: 5271: 5265: 5263: 5257: 5256: 5251: 5250: 5243: 5236: 5228: 5219: 5218: 5216: 5215: 5203: 5190: 5187: 5186: 5183: 5182: 5180: 5179: 5174: 5169: 5164: 5159: 5157:Paul R. McHugh 5154: 5148: 5146: 5142: 5141: 5139: 5138: 5133: 5128: 5123: 5117: 5115: 5111: 5110: 5108: 5107: 5102: 5097: 5092: 5087: 5082: 5077: 5072: 5067: 5062: 5057: 5052: 5047: 5042: 5037: 5032: 5027: 5022: 5017: 5015:Ivan Izquierdo 5012: 5007: 5002: 4997: 4992: 4987: 4982: 4976: 4974: 4967: 4963: 4962: 4960: 4959: 4952: 4942: 4941: 4940: 4930: 4925: 4920: 4915: 4914: 4913: 4903: 4898: 4893: 4888: 4883: 4873: 4868: 4862: 4860: 4856: 4855: 4853: 4852: 4847: 4846: 4845: 4834: 4832: 4828: 4827: 4825: 4824: 4819: 4814: 4809: 4804: 4799: 4794: 4789: 4788: 4787: 4782: 4772: 4766: 4764: 4760: 4759: 4757: 4756: 4751: 4746: 4741: 4736: 4735: 4734: 4723: 4721: 4717: 4716: 4714: 4713: 4708: 4703: 4697: 4695: 4689: 4688: 4686: 4685: 4680: 4679: 4678: 4668: 4663: 4657: 4655: 4649: 4648: 4646: 4645: 4640: 4635: 4630: 4625: 4620: 4615: 4613:Hindsight bias 4610: 4605: 4599: 4597: 4591: 4590: 4588: 4587: 4582: 4577: 4572: 4567: 4562: 4560:Memory erasure 4557: 4552: 4547: 4542: 4541: 4540: 4535: 4530: 4525: 4520: 4518:post-traumatic 4515: 4510: 4505: 4494: 4492: 4486: 4485: 4482: 4481: 4479: 4478: 4473: 4468: 4463: 4458: 4456:Personal-event 4453: 4448: 4443: 4438: 4433: 4432: 4431: 4426: 4421: 4411: 4406: 4400: 4398: 4392: 4391: 4389: 4388: 4386:Working memory 4383: 4375: 4373: 4367: 4366: 4364: 4363: 4358: 4356:Motor learning 4353: 4348: 4343: 4338: 4333: 4327: 4325: 4316: 4312: 4311: 4309: 4308: 4303: 4298: 4292: 4291: 4286: 4281: 4275: 4273: 4272:Basic concepts 4269: 4268: 4261: 4260: 4253: 4246: 4238: 4232: 4231: 4202:10.1.1.90.9696 4184: 4147: 4121:(9): 456–461. 4110: 4074:(9): 850–855. 4059: 4004: 3976:(2): 147–153. 3959: 3956: 3954: 3953: 3928: 3904: 3879: 3852:(2): 317–330. 3823: 3796: 3730: 3675: 3664:(2): 109–136. 3636: 3613: 3586:(2): 123–132. 3568: 3539:(4): 695–701. 3519: 3462: 3403: 3360: 3311: 3262: 3203: 3138: 3087: 3043: 2992: 2949: 2874: 2849: 2790: 2753:(9): 850–855. 2728: 2693:(5): 793–802. 2677: 2623: 2594:(1): 135–148. 2563: 2534:(5): 500–510. 2509: 2472:(4): 911–917. 2453: 2432:(3): 377–385. 2409: 2356: 2303: 2296: 2262: 2243:(2): 145–153. 2227: 2200:(2): 205–210. 2181: 2129: 2079: 2052:(2): 217–227. 2013: 1986:(2): 119–130. 1967: 1940:(2): 169–177. 1919: 1912: 1889: 1860:(3): 201–211. 1837: 1830: 1807: 1750: 1712: 1691:(9): 711–716. 1671: 1611: 1590:(2): 238–252. 1567: 1540:(4): 262–275. 1505: 1462: 1418: 1407:(3): 215–234. 1391: 1337: 1283: 1239: 1183: 1137: 1082: 1080: 1077: 1076: 1075: 1070: 1065: 1060: 1055: 1048: 1045: 1031: 1028: 961: 958: 883: 880: 870:, and various 778: 775: 742: 739: 715: 712: 708:sleep spindles 699: 696: 626: 623: 581: 578: 555: 552: 534: 531: 525: 522: 499: 496: 428: 427:Standard model 425: 403: 400: 372: 369: 347:Spacing effect 345:Main article: 342: 341:Spacing effect 339: 284:Main article: 281: 278: 249: 248:Standard model 246: 238:Late-phase LTP 213: 210: 187:preparations, 112:Henry Molaison 64: 61: 15: 9: 6: 4: 3: 2: 5587: 5576: 5573: 5571: 5568: 5567: 5565: 5550: 5547: 5545: 5542: 5540: 5537: 5535: 5532: 5530: 5527: 5525: 5522: 5520: 5517: 5515: 5512: 5510: 5507: 5505: 5502: 5500: 5497: 5495: 5492: 5491: 5489: 5485: 5479: 5476: 5474: 5471: 5469: 5466: 5464: 5463:Consolidation 5461: 5460: 5458: 5456: 5452: 5446: 5443: 5439: 5436: 5434: 5431: 5429: 5426: 5425: 5424: 5421: 5419: 5416: 5414: 5411: 5409: 5406: 5403: 5399: 5396: 5394: 5391: 5389: 5386: 5382: 5379: 5378: 5377: 5374: 5372: 5369: 5368: 5366: 5364: 5360: 5354: 5351: 5347: 5344: 5342: 5339: 5338: 5337: 5334: 5332: 5329: 5327: 5324: 5322: 5319: 5317: 5314: 5312: 5311:Consciousness 5309: 5307: 5306:Comprehension 5304: 5302: 5299: 5295: 5292: 5291: 5290: 5287: 5285: 5282: 5280: 5277: 5275: 5272: 5270: 5267: 5266: 5264: 5262: 5258: 5249: 5244: 5242: 5237: 5235: 5230: 5229: 5226: 5214: 5204: 5202: 5192: 5191: 5188: 5178: 5175: 5173: 5170: 5168: 5165: 5163: 5160: 5158: 5155: 5153: 5150: 5149: 5147: 5143: 5137: 5136:Clive Wearing 5134: 5132: 5129: 5127: 5124: 5122: 5119: 5118: 5116: 5112: 5106: 5103: 5101: 5100:Endel Tulving 5098: 5096: 5095:Anne Treisman 5093: 5091: 5088: 5086: 5083: 5081: 5078: 5076: 5073: 5071: 5068: 5066: 5063: 5061: 5058: 5056: 5055:Brenda Milner 5053: 5051: 5048: 5046: 5043: 5041: 5040:James McGaugh 5038: 5036: 5033: 5031: 5028: 5026: 5023: 5021: 5018: 5016: 5013: 5011: 5008: 5006: 5005:Sigmund Freud 5003: 5001: 4998: 4996: 4993: 4991: 4988: 4986: 4983: 4981: 4978: 4977: 4975: 4971: 4968: 4964: 4958: 4957: 4953: 4950: 4949:retrospective 4946: 4943: 4939: 4936: 4935: 4934: 4931: 4929: 4928:Muscle memory 4926: 4924: 4921: 4919: 4916: 4912: 4909: 4908: 4907: 4904: 4902: 4899: 4897: 4894: 4892: 4889: 4887: 4884: 4881: 4877: 4874: 4872: 4869: 4867: 4864: 4863: 4861: 4857: 4851: 4848: 4844: 4841: 4840: 4839: 4836: 4835: 4833: 4829: 4823: 4820: 4818: 4815: 4813: 4810: 4808: 4805: 4803: 4800: 4798: 4795: 4793: 4790: 4786: 4783: 4781: 4778: 4777: 4776: 4775:Art of memory 4773: 4771: 4768: 4767: 4765: 4761: 4755: 4752: 4750: 4747: 4745: 4742: 4740: 4737: 4733: 4730: 4729: 4728: 4725: 4724: 4722: 4718: 4712: 4709: 4707: 4704: 4702: 4699: 4698: 4696: 4694: 4690: 4684: 4681: 4677: 4674: 4673: 4672: 4669: 4667: 4664: 4662: 4659: 4658: 4656: 4654: 4650: 4644: 4641: 4639: 4636: 4634: 4631: 4629: 4626: 4624: 4623:Memory biases 4621: 4619: 4616: 4614: 4611: 4609: 4606: 4604: 4603:Confabulation 4601: 4600: 4598: 4596: 4595:Memory errors 4592: 4586: 4583: 4581: 4578: 4576: 4573: 4571: 4568: 4566: 4563: 4561: 4558: 4556: 4553: 4551: 4548: 4546: 4543: 4539: 4536: 4534: 4531: 4529: 4526: 4524: 4521: 4519: 4516: 4514: 4513:post-hypnotic 4511: 4509: 4506: 4504: 4501: 4500: 4499: 4496: 4495: 4493: 4491: 4487: 4477: 4474: 4472: 4469: 4467: 4466:Rote learning 4464: 4462: 4459: 4457: 4454: 4452: 4449: 4447: 4444: 4442: 4441:Hyperthymesia 4439: 4437: 4434: 4430: 4427: 4425: 4422: 4420: 4417: 4416: 4415: 4412: 4410: 4407: 4405: 4404:Active recall 4402: 4401: 4399: 4397: 4393: 4387: 4384: 4381: 4377: 4376: 4374: 4372: 4368: 4362: 4359: 4357: 4354: 4352: 4349: 4347: 4344: 4342: 4339: 4337: 4334: 4332: 4329: 4328: 4326: 4324: 4320: 4317: 4313: 4307: 4304: 4302: 4301:Consolidation 4299: 4297: 4294: 4293: 4290: 4287: 4285: 4282: 4280: 4277: 4276: 4274: 4270: 4266: 4259: 4254: 4252: 4247: 4245: 4240: 4239: 4236: 4228: 4224: 4220: 4216: 4212: 4208: 4203: 4198: 4195:(4): 352–68. 4194: 4190: 4185: 4181: 4177: 4173: 4169: 4165: 4161: 4157: 4153: 4148: 4144: 4140: 4136: 4132: 4128: 4124: 4120: 4116: 4111: 4107: 4103: 4098: 4093: 4089: 4085: 4081: 4077: 4073: 4069: 4065: 4060: 4056: 4052: 4048: 4044: 4040: 4036: 4031: 4026: 4022: 4018: 4014: 4010: 4005: 4001: 3997: 3992: 3987: 3983: 3979: 3975: 3971: 3967: 3962: 3961: 3942: 3938: 3932: 3924: 3920: 3919: 3914: 3908: 3893: 3889: 3883: 3875: 3871: 3867: 3863: 3859: 3855: 3851: 3847: 3846: 3841: 3834: 3830: 3826: 3824:9780415897167 3820: 3816: 3812: 3811: 3803: 3801: 3792: 3788: 3783: 3778: 3774: 3770: 3766: 3762: 3758: 3754: 3753: 3748: 3744: 3740: 3734: 3726: 3722: 3718: 3714: 3710: 3706: 3705: 3697: 3693: 3689: 3682: 3680: 3671: 3667: 3663: 3659: 3658: 3650: 3643: 3641: 3632: 3628: 3624: 3617: 3609: 3605: 3601: 3597: 3593: 3589: 3585: 3581: 3580: 3572: 3564: 3560: 3556: 3552: 3547: 3542: 3538: 3534: 3530: 3523: 3515: 3511: 3507: 3503: 3498: 3493: 3489: 3485: 3481: 3477: 3473: 3466: 3458: 3454: 3449: 3444: 3439: 3434: 3430: 3426: 3422: 3418: 3414: 3407: 3399: 3395: 3391: 3387: 3383: 3379: 3375: 3371: 3364: 3356: 3352: 3347: 3342: 3338: 3334: 3330: 3326: 3322: 3315: 3307: 3303: 3298: 3293: 3289: 3285: 3281: 3277: 3273: 3266: 3258: 3254: 3249: 3244: 3239: 3234: 3230: 3226: 3223:(2): e30502. 3222: 3218: 3214: 3207: 3199: 3195: 3190: 3185: 3181: 3177: 3173: 3169: 3165: 3161: 3157: 3153: 3149: 3142: 3134: 3130: 3126: 3122: 3118: 3114: 3110: 3106: 3102: 3098: 3091: 3083: 3079: 3075: 3071: 3067: 3063: 3059: 3055: 3047: 3039: 3035: 3031: 3027: 3023: 3019: 3015: 3011: 3007: 3003: 2996: 2988: 2984: 2980: 2976: 2972: 2968: 2964: 2960: 2953: 2946: 2941: 2935: 2931: 2926: 2921: 2917: 2913: 2909: 2905: 2901: 2897: 2893: 2889: 2885: 2878: 2863: 2859: 2853: 2845: 2841: 2836: 2831: 2827: 2823: 2818: 2813: 2809: 2805: 2801: 2794: 2786: 2782: 2777: 2772: 2768: 2764: 2760: 2756: 2752: 2748: 2744: 2737: 2735: 2733: 2724: 2720: 2716: 2712: 2708: 2704: 2700: 2696: 2692: 2688: 2681: 2673: 2669: 2665: 2661: 2657: 2653: 2649: 2645: 2641: 2637: 2630: 2628: 2619: 2615: 2611: 2607: 2602: 2597: 2593: 2589: 2585: 2578: 2576: 2574: 2572: 2570: 2568: 2559: 2555: 2550: 2545: 2541: 2537: 2533: 2529: 2525: 2518: 2516: 2514: 2505: 2501: 2497: 2493: 2489: 2485: 2480: 2475: 2471: 2467: 2460: 2458: 2449: 2445: 2440: 2435: 2431: 2427: 2426:Psychobiology 2423: 2416: 2414: 2405: 2401: 2397: 2393: 2389: 2385: 2382:(2): 225–33. 2381: 2377: 2370: 2363: 2361: 2352: 2348: 2344: 2340: 2336: 2332: 2329:(1): 125–33. 2328: 2324: 2317: 2310: 2308: 2299: 2293: 2289: 2285: 2281: 2277: 2273: 2266: 2258: 2254: 2250: 2246: 2242: 2238: 2231: 2223: 2219: 2215: 2211: 2207: 2203: 2199: 2195: 2188: 2186: 2177: 2173: 2169: 2165: 2161: 2157: 2153: 2149: 2142: 2140: 2138: 2136: 2134: 2125: 2121: 2117: 2113: 2109: 2108:10.1038/nn739 2105: 2101: 2097: 2090: 2088: 2086: 2084: 2075: 2071: 2067: 2063: 2059: 2055: 2051: 2047: 2040: 2038: 2036: 2034: 2032: 2030: 2028: 2026: 2024: 2022: 2020: 2018: 2009: 2005: 2001: 1997: 1993: 1989: 1985: 1981: 1974: 1972: 1963: 1959: 1955: 1951: 1947: 1943: 1939: 1935: 1928: 1926: 1924: 1915: 1909: 1905: 1904: 1896: 1894: 1885: 1881: 1876: 1871: 1867: 1863: 1859: 1855: 1851: 1844: 1842: 1833: 1827: 1823: 1822: 1814: 1812: 1803: 1799: 1795: 1791: 1787: 1783: 1778: 1773: 1769: 1765: 1761: 1754: 1746: 1742: 1738: 1734: 1730: 1726: 1719: 1717: 1708: 1704: 1699: 1694: 1690: 1686: 1682: 1675: 1667: 1663: 1659: 1655: 1650: 1649:2027.42/56238 1645: 1641: 1637: 1633: 1629: 1622: 1615: 1607: 1603: 1598: 1593: 1589: 1585: 1581: 1577: 1571: 1563: 1559: 1555: 1551: 1547: 1543: 1539: 1535: 1528: 1526: 1524: 1522: 1520: 1518: 1516: 1514: 1512: 1510: 1501: 1497: 1493: 1489: 1485: 1481: 1477: 1473: 1466: 1458: 1454: 1449: 1444: 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