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Long-term memory

Long-term memory (LTM) is the brain's capacity to store, retain, and retrieve information and experiences over extended periods, potentially lasting a lifetime, distinguishing it from shorter-duration forms like sensory or working memory.^[1] This system enables individuals to learn from past events, form personal identity, and navigate complex environments by encoding knowledge that shapes behavior and decision-making.^[1] In the classic Atkinson-Shiffrin model of human memory, proposed in 1968, LTM serves as the third and most enduring stage, following sensory memory and short-term (or working) memory, where information is transferred through rehearsal and consolidation for indefinite storage.^[2] LTM is broadly categorized into two main types: declarative memory, which involves consciously accessible facts and events, and procedural memory, which encompasses unconscious skills and habits acquired through repetition.^[1] Declarative memory further divides into episodic memory for personal, time-stamped experiences (e.g., recalling a specific birthday celebration) and semantic memory for general knowledge and facts (e.g., knowing the capital of France).^[1] Procedural memory, in contrast, supports automated actions like riding a bicycle or playing an instrument, often without deliberate recall.^[1] The formation of LTM relies on synaptic plasticity, particularly long-term potentiation (LTP), a process discovered in the 1960s that strengthens neural connections between neurons, allowing memories to persist beyond initial exposure.^[1] Consolidation typically begins in the hippocampus, which temporarily holds and organizes new memories before distributing them across the cerebral cortex for long-term storage, a mechanism involving protein synthesis and gene expression.^[1] Key brain regions include the amygdala for tagging emotional significance to memories, enhancing their retention, and the cerebellum and basal ganglia for procedural learning.^[1] Retrieval from LTM can be effortful (e.g., searching for a forgotten name) or automatic (e.g., recognizing a familiar face), influenced by cues, context, and interference from similar memories.^[1] Disruptions in LTM, such as those seen in Alzheimer's disease or amnesia, highlight its vulnerability to neurological damage, underscoring its role in cognitive health.^[1]

Overview

Definition and Characteristics

Long-term memory (LTM) refers to the brain's system for storing information and experiences over extended periods, often indefinitely, serving as a repository for knowledge that persists beyond immediate use.^[1] This contrasts with temporary storage mechanisms, where information is held only briefly for processing.^[3] In cognitive psychology, LTM is conceptualized as the final stage in memory formation, where selected information from earlier processing stages is transferred and maintained for potential later retrieval.^[4] Key characteristics of long-term memory include its virtually unlimited capacity, allowing storage of vast amounts of data without apparent overflow, and durations that range from days, months, or years to an entire lifetime.^[5] Unlike fleeting memories, LTM exhibits resistance to decay, with encoded information remaining stable over time unless disrupted by factors such as interference or neurological damage.^[6] This stability enables the retention of complex structures, such as semantic networks of facts or procedural routines, supporting adaptive behavior across contexts.^[7] Everyday examples of long-term memories illustrate these traits, including autobiographical recollections like the details of one's first school day or semantic knowledge such as historical events learned in childhood.^[8] Procedural memories, such as riding a bicycle or playing a musical instrument acquired years prior, also exemplify LTM's enduring nature and resistance to loss.^[9] The concept of long-term memory was initially formalized in the 1960s through the multi-store model proposed by psychologists Richard C. Atkinson and Richard M. Shiffrin, who described it as a permanent store distinct from sensory and short-term registers.^[3] This framework, introduced in their 1968 paper, laid the groundwork for subsequent research by emphasizing LTM's role in long-lasting retention.^[3]

Distinction from Short-Term Memory

Long-term memory (LTM) differs from short-term memory (STM) primarily in terms of capacity and duration of storage. STM has a limited capacity of approximately 7 ± 2 items or chunks, as demonstrated by Miller's (1956) analysis of immediate memory span across various tasks like digit recall.^[10] In contrast, LTM exhibits a vast, potentially unlimited capacity, with evidence from visual memory studies showing individuals can store and retrieve details of at least 2,500 unique objects across numerous scenes with high accuracy.^[11] Duration in STM is brief, typically lasting 15 to 30 seconds without active maintenance, while LTM retains information for years or a lifetime, allowing persistent access to encoded experiences.^[2] Neuropsychological evidence underscores this separation, particularly through cases of anterograde amnesia where STM remains functional but new LTM formation is impaired. The landmark case of patient H.M., who underwent bilateral medial temporal lobe resection in 1953, preserved a normal digit span of seven items in immediate recall tasks but could not consolidate new episodic memories beyond a few minutes, revealing a profound dissociation between the systems.^[12] Such findings indicate that while STM supports temporary holding, LTM requires distinct mechanisms for enduring storage. Behavioral experiments further highlight these differences via the serial position effect observed in list-learning tasks. When participants recall a sequence of words, early items benefit from a primacy effect due to deeper encoding into LTM through extended rehearsal, whereas later items show a recency effect attributable to fresh retention in STM.^[13] Introducing a distractor task, such as counting backward, immediately after presentation disrupts the recency effect by preventing reliance on STM but leaves the primacy effect intact, confirming independent contributions from LTM.^[13] The transfer of information from STM to LTM occurs primarily through rehearsal processes that reinforce memory traces for consolidation. Maintenance rehearsal, involving repetition of items, extends STM duration and facilitates encoding into LTM, as conceptualized in the multi-store model where such mechanisms bridge the two stores.^[2]

Theoretical Models

Dual-Store Model

The dual-store model, also known as the multi-store model, posits that human memory operates through distinct stages, with long-term memory (LTM) serving as the primary repository for enduring information storage.^[2] Proposed by Atkinson and Shiffrin in 1968, the model describes a sequential flow beginning with sensory memory, a brief register of environmental stimuli lasting fractions of a second to seconds, which transfers attended information to a short-term store (STS) of limited capacity (approximately 7 ± 2 items) and duration (about 20-30 seconds without rehearsal).^[2] From STS, information enters LTM through rehearsal processes: maintenance rehearsal sustains items temporarily in STS, while elaborative rehearsal promotes deeper semantic processing for more permanent transfer to LTM, conceived as an unlimited-capacity, long-lasting archive.^[2] This architecture emphasizes control processes, such as attention and retrieval strategies, that govern the transition between stores.^[2] A key feature of LTM in the dual-store framework is its role as a stable, context-dependent repository, where retrieval success depends on the match between encoding and retrieval conditions, as articulated in Tulving's encoding specificity principle from the 1970s.^[14] This principle holds that cues effective for accessing LTM traces are those present or similar to those during initial encoding, underscoring LTM's reliance on associative networks rather than mere strength of storage. Building on the Atkinson-Shiffrin model, Baddeley extended the conceptualization of STS in the 1970s by reframing it as working memory (WM), a dynamic system interacting bidirectionally with LTM.^[15] Baddeley's 1974 model introduces a central executive for attentional control and coordination, a phonological loop for verbal-auditory information, and a visuospatial sketchpad for visual-spatial material, enabling WM to manipulate and integrate LTM contents for tasks like reasoning or comprehension.^[15] Empirical support for the model's distinction between STS and LTM comes from free recall experiments, where performance over long delays (e.g., minutes to days) reveals LTM's dominance, as short-term effects like the recency portion of the serial position curve diminish while primacy effects—attributable to LTM rehearsal—persist.^[2] In such studies, participants recalling word lists after extended intervals show recall probabilities approaching those predicted by LTM transfer rates, with rehearsal manipulations enhancing long-delay performance beyond what STS alone could sustain.^[2] These findings highlight the model's utility in explaining how information consolidates into LTM, though later refinements addressed WM's active role in modulating LTM access.^[15]

Single-Store Model

The single-store model, also known as the unitary store hypothesis, posits that memory operates as a single system where all information is stored in one repository, varying only in trace strength along a continuum rather than being divided into distinct short-term and long-term stores.^[16] This view, articulated early by Arthur W. Melton, argues that apparent differences between short- and long-term memory arise from factors like encoding depth, retrieval cues, and interference rather than separate structural components.^[17] In contrast to the dual-store model of Atkinson and Shiffrin, which posits a sharp boundary between transient short-term storage and permanent long-term storage, the single-store approach suggests a more integrated process without such categorical divisions.^[16] A key proponent of this perspective in the 1970s was Bennet B. Murdock, who developed a trace decay model emphasizing that memory traces weaken gradually over time based on factors like rehearsal and interference, forming a continuum of accessibility without discrete store boundaries.^[18] Murdock's framework, detailed in his 1974 book Human Memory: Theory and Data, treats forgetting as a unified process where trace strength diminishes according to power-law functions, challenging the idea of rapid decay confined to a short-term store followed by stable long-term retention.^[18] Supporting evidence includes the observation of smooth, gradient forgetting curves across time scales, which follow a power-law decline in recall probability rather than exhibiting the abrupt shifts predicted by dual-store theories.^[19] For instance, studies on verbal learning show that retention decreases continuously from seconds to days, with no clear inflection point demarcating short- from long-term phases.^[20] Further contradictory evidence to the dual-store model comes from interference paradigms demonstrating effects that span what would be separate stores. Proactive and retroactive interference occur across varying retention intervals, with similar items from long-past learning disrupting immediate recall, indicating a shared underlying mechanism rather than isolated stores.^[21] Key studies, such as those using paired-associate learning, reveal no sharp cutoff in susceptibility to interference between short and extended delays, as intrusions from prior lists affect performance uniformly along the memory continuum.^[22] Modern variants of the single-store model incorporate distributed representations, where long-term memory emerges from repeated activations in a unified network without a dedicated short-term phase. Murdock's later Theory of Distributed Associative Memory (TODAM), refined in the 1990s, models memory as vector-based traces stored via convolution in a single associative space, accounting for item, order, and associative information through probabilistic retrieval.^[23] Empirical support includes recency effects in free-recall tasks under continuous distractors, where enhanced recall of recent items persists even with interpolated activity, suggesting temporal context gradients within one store rather than reliance on a fragile short-term buffer.^[24] Computational simulations of these paradigms, such as those revisiting short-term memory's role in recency, reinforce the single-store view by fitting data with strength-based activation alone.^[25]

Divisions

Explicit Memory

Explicit memory, also known as declarative memory, is the conscious, intentional recollection of factual information, previous experiences, and concepts that can be verbally expressed or communicated.^[26] This form of long-term memory enables individuals to deliberately access and articulate stored knowledge, distinguishing it from unconscious memory processes. Endel Tulving conceptualized explicit memory as part of a declarative system, emphasizing its role in representing information that is accessible through awareness and reflection.^[27] Explicit memory is primarily divided into two subtypes: episodic and semantic. Episodic memory involves the vivid recollection of personal events situated in specific times and contexts, often accompanied by a sense of subjective time travel, such as remembering the details of a childhood birthday celebration.^[26] In contrast, semantic memory encompasses general factual knowledge detached from personal context, including concepts like the capital of France being Paris or the basic principles of mathematics.^[26] Autobiographical memory represents an integration of these subtypes, forming a cohesive narrative of one's life experiences by combining episodic details with semantic understanding of the self.^[28] These components support essential cognitive functions, including learning through the accumulation and conscious retrieval of information, and decision-making by providing factual bases for evaluating options and anticipating outcomes.^[29] Explicit memory is commonly evaluated using recall tasks, where individuals reproduce information from memory, or recognition tasks, where they identify previously encountered items among distractors.^[30] The development of explicit memory begins in early childhood, emerging around age 2 to 3 years alongside language acquisition and the maturation of brain structures like the hippocampus, with significant improvements continuing through adolescence to peak in early adulthood.^[26] Unlike implicit memory, which influences behavior without awareness, explicit memory relies on conscious effort, allowing for reflective and strategic use in complex cognition.^[26]

Implicit Memory

Implicit memory, also referred to as nondeclarative memory, is a category of long-term memory that operates unconsciously to influence thoughts, perceptions, and behaviors without requiring deliberate recollection or awareness of past experiences.^[31] Unlike explicit memory, it manifests through facilitated performance on tasks, such as improved reaction times or automatic skill execution, rather than through verbal reports of remembering.^[31] For instance, priming effects exemplify this, where prior exposure to a stimulus enhances subsequent processing of related information, like completing word fragments more quickly after seeing related cues.^[32] This form of memory includes distinct subtypes: procedural memory, priming, and conditioning. Procedural memory encompasses the learning and retention of motor skills and cognitive routines, enabling automatic execution of complex actions such as riding a bicycle or playing a musical instrument once mastered through repetition.^[33] Priming involves subtle perceptual or conceptual enhancements from previous encounters, leading to faster or more accurate identification without conscious retrieval of the initial exposure.^[32] Conditioning, including classical and operant forms, establishes unconscious associations between stimuli and responses, such as an automatic emotional reaction to a previously neutral cue paired with an aversive event.^[34] Compelling evidence for implicit memory's independence from conscious processes emerges from studies of amnesic patients, including the landmark case of H.M. (Henry Molaison), who suffered profound hippocampal damage resulting in anterograde amnesia. Despite inability to recall training sessions, H.M. progressively improved at mirror-drawing tasks over multiple trials, demonstrating retention comparable to healthy individuals.^[35] Similar preservation occurs in other amnesics for procedural skills and classical conditioning paradigms, where they acquire conditioned responses without episodic memory of the pairings.^[33]^[34] These findings indicate that implicit memory relies on brain systems distinct from those supporting explicit recall, notably functioning without the hippocampus.^[31] In everyday cognition, implicit memory facilitates habit formation by forging context-response associations through repetition, allowing behaviors like teeth brushing to become effortless and cue-driven over time.^[36] It also drives expertise development, as repeated practice transforms deliberate actions into fluid, unconscious competencies, evident in skilled performers who execute routines with minimal cognitive effort.^[33]

Encoding and Consolidation

Encoding Processes

Encoding refers to the initial processes by which information from sensory input or working memory is transformed and stored in long-term memory, determining the durability and accessibility of memories. This stage involves active cognitive operations that convert transient experiences into lasting representations, often requiring transfer from short-term storage mechanisms as described in multi-store models.^[37] A foundational framework for understanding encoding is the levels of processing theory, which posits that the depth of analysis applied to information influences its retention in long-term memory. Shallow processing focuses on superficial features, such as the physical or sensory attributes of a stimulus (e.g., the font or color of a word), leading to weaker memory traces. In contrast, deeper semantic processing involves meaningful interpretation, like relating the word to its concept or personal associations, resulting in more robust encoding and better recall. This graded effect was empirically demonstrated through experiments showing superior recognition for semantically processed items compared to those processed phonetically or structurally.^[37] Several factors modulate the effectiveness of encoding. Attention serves as a critical gatekeeper, selectively directing resources to relevant stimuli and facilitating deeper processing; divided attention during encoding impairs the formation of durable long-term memories by limiting the elaboration of representations. Emotional arousal enhances encoding via amygdala-mediated mechanisms, which amplify consolidation for affectively charged events, such as fearful or rewarding experiences, leading to prioritized storage of survival-relevant information. Repetition strengthens encoding, but the spacing effect reveals that distributed practice—revisiting material over increasing intervals—is more effective for long-term retention than massed repetition, as it promotes varied contextual cues and reduces forgetting.^[38]^[39]^[40] Intentional strategies can optimize encoding by imposing structure on information. Mnemonics, such as the method of loci (associating items with familiar spatial locations) or pegword systems (linking new data to pre-learned rhymes), leverage imagery and associations to create elaborate, retrievable traces that outperform rote memorization for lists or sequences. Chunking reorganizes information into meaningful units, expanding effective capacity beyond isolated items; for instance, grouping digits into phone numbers allows recall of larger amounts as cohesive patterns rather than discrete elements.^[41]^[42] Integrating multiple sensory modalities during encoding fosters stronger memory traces by creating richer, interconnected representations. Multisensory experiences, like combining visual and auditory inputs, enhance perceptual salience and discrimination, leading to improved long-term recognition compared to unisensory learning, as the brain binds cross-modal cues into unified episodes.^[43]

Role of Sleep and Consolidation

Sleep plays a crucial role in the consolidation of memories, transforming initially fragile traces into stable long-term representations through offline processing that occurs primarily during non-wakeful states. This process, known as memory consolidation, involves two main types: synaptic consolidation, which strengthens local neural connections at the site of encoding to make memories more resistant to interference, and systems consolidation, which reorganizes memory traces across brain networks, gradually transferring dependence from temporary storage to distributed cortical sites for long-term retention. Synaptic consolidation occurs rapidly, often within hours, while systems consolidation unfolds over days to years, with sleep facilitating both by providing a period of reduced sensory input and enhanced neural replay.^[44]^[45]^[46] Different sleep stages contribute distinctly to consolidation, with slow-wave sleep (SWS), characterized by high-amplitude delta oscillations, being particularly important for declarative memories such as facts and events. During SWS, the neocortex and hippocampus engage in a dialogue that replays and integrates new information, enhancing the stability of hippocampal-dependent memories through coordinated slow oscillations and sleep spindles. In contrast, rapid eye movement (REM) sleep supports the consolidation of procedural skills, like motor sequences, and emotional memories, potentially by facilitating the integration of affective elements into broader networks via theta activity and heightened acetylcholine levels.^[47]^[48]^[49]^[50] Pioneering evidence for sleep's role comes from animal studies demonstrating memory replay, where neural patterns active during wakeful learning are reactivated during sleep. In a seminal experiment, recordings from rat hippocampal place cells showed that sequences of cell firing experienced during spatial navigation were replayed in the same order during subsequent SWS, suggesting a mechanism for strengthening spatial memories offline. Human studies further corroborate this, revealing that sleep deprivation after learning impairs long-term memory performance; for instance, one night without sleep reduces recall of declarative material by up to 40% compared to rested controls, highlighting sleep's necessity for effective consolidation.^[51]^[52]^[53]^[54] Recent advances in the 2020s have utilized targeted memory reactivation (TMR), a technique where sensory cues associated with learning are re-presented during sleep to enhance specific memories, with neuroimaging providing mechanistic insights. Functional MRI studies during TMR in SWS and REM have shown increased reactivation of task-related brain patterns, correlating with improved memory accuracy upon waking; for example, odor or auditory cues during sleep can enhance consolidation of emotional memories, as evidenced by strengthened hippocampal-cortical connectivity. These findings underscore TMR's potential to selectively stabilize long-term memories, building on natural replay processes observed in both animals and humans.^[55]^[56]^[57]

Neural and Biological Basis

Brain Structures and Pathways

The hippocampus, located within the medial temporal lobe, plays a central role in the initial encoding and consolidation of long-term memories, particularly declarative ones such as episodic and semantic information.^[35] Damage to the hippocampus, as seen in the case of patient H.M. who underwent bilateral removal in 1953, results in anterograde amnesia, preventing the formation of new explicit long-term memories while sparing pre-existing ones and implicit memory functions.^[35] The medial temporal lobe, encompassing the hippocampus, entorhinal cortex, and perirhinal cortex, supports episodic memory by integrating contextual details from experiences into coherent representations.^[58] The prefrontal cortex contributes to semantic memory organization and retrieval processes, facilitating the strategic access and manipulation of stored knowledge across distributed networks.^[59] Long-term memories are ultimately stored in a distributed manner across the neocortex, where sensory-specific regions like the temporal and parietal lobes maintain perceptual and associative components after consolidation.^[60] The amygdala, situated in the medial temporal lobe as part of the limbic system, modulates the consolidation and persistence of emotionally salient long-term memories by interacting with the hippocampus and other regions, enhancing retention through noradrenergic and stress hormone signaling.^[61] Key neural pathways underpin these functions, with the Papez circuit serving as a critical loop for memory processing: it connects the hippocampus via the fornix to the mammillary bodies, then to the anterior thalamus, cingulate gyrus, and back to the entorhinal cortex, enabling the recirculation and stabilization of memory traces.^[62] This circuit, originally proposed for emotional processing, has been established as essential for long-term memory formation and retrieval, with disruptions leading to deficits in episodic recall.^[63] For implicit memories, such as procedural skills, the basal ganglia form parallel pathways involving the striatum and substantia nigra, supporting habit formation and non-declarative learning independent of conscious awareness.^[64] The cerebellum contributes to procedural long-term memory, particularly for motor skills and sequence learning, by refining timing and coordination through its Purkinje cells and mossy fiber-granule cell circuits, enabling the automation of practiced actions.^[65] Functional imaging studies provide evidence for these structures and pathways through activation patterns during memory tasks. Positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) reveal heightened hippocampal and medial temporal lobe activity during encoding of novel information, with subsequent neocortical engagement during retrieval, confirming the transition from temporary to permanent storage.^[66] Lesion studies, including H.M.'s, corroborate this by showing that isolated hippocampal damage impairs consolidation without affecting remote memories already distributed to the neocortex.^[35] Lateralization further refines these processes, with the left hemisphere, particularly the left hippocampus and prefrontal regions, specializing in verbal and semantic memory, while the right hemisphere handles spatial and visuospatial components.^[67] This hemispheric asymmetry is evident in fMRI activations during verbal versus spatial tasks, highlighting adaptive specialization in long-term memory networks.^[68]

Cellular and Molecular Mechanisms

Long-term potentiation (LTP) represents a fundamental form of synaptic plasticity implicated in the cellular basis of long-term memory storage. Initially discovered in the hippocampus, LTP involves a persistent strengthening of synaptic efficacy following high-frequency stimulation of afferent pathways. This process is critically dependent on N-methyl-D-aspartate (NMDA) receptors, which, upon activation by coincident presynaptic glutamate release and postsynaptic depolarization, permit calcium influx that triggers intracellular signaling cascades leading to synaptic enhancement.^[69] The principle of Hebbian learning underpins LTP, positing that synaptic connections between neurons strengthen when presynaptic activity repeatedly precedes postsynaptic firing, famously summarized as "cells that fire together wire together." This mechanism aligns with the associative nature of memory formation, where correlated neural activity stabilizes synaptic weights to encode information. Experimental evidence from hippocampal slices demonstrates that blocking NMDA receptor function prevents LTP induction, underscoring its necessity for Hebbian plasticity.^[69] At the molecular level, the transition from early-phase LTP (E-LTP), which is transient and protein synthesis-independent, to late-phase LTP (L-LTP), lasting hours to days, requires de novo gene expression and protein synthesis. Key players include the immediate early gene Arc (activity-regulated cytoskeleton-associated protein), which is rapidly transcribed in response to synaptic activity and regulates AMPA receptor trafficking to stabilize potentiated synapses. Similarly, brain-derived neurotrophic factor (BDNF) promotes dendritic spine morphogenesis and synaptic consolidation by activating TrkB receptors, facilitating local protein synthesis essential for L-LTP maintenance. Inhibition of protein synthesis, such as with anisomycin, abolishes L-LTP while sparing E-LTP, confirming its role in enduring synaptic changes underlying long-term memory.^[70] Epigenetic modifications, particularly DNA methylation and demethylation, provide stable yet dynamic tags that influence gene expression for memory stabilization. In fear memory paradigms, increased DNA methylation at promoter regions of plasticity-related genes, such as BDNF, represses transcription during consolidation, while active demethylation via TET enzymes enables rapid reactivation. Studies from the 2010s using contextual fear conditioning in rodents revealed that intra-hippocampal infusion of DNA methyltransferase inhibitors impairs fear memory formation, whereas demethylating agents enhance extinction, highlighting bidirectional epigenetic control in memory processes. These modifications persist beyond initial encoding, contributing to the long-term retention of emotional memories.^[71]^[72] Memory engrams, conceptualized as physical traces encoded in sparse ensembles of neurons, have been directly visualized and manipulated using optogenetic techniques. Pioneering work in the 2010s demonstrated that engram cells in the hippocampus, labeled by activity-dependent expression of Channelrhodopsin-2 during fear learning, can be reactivated with light to elicit memory recall or even implant false memories. These engram ensembles exhibit heightened synaptic plasticity markers, including elevated Arc and BDNF expression, integrating cellular mechanisms into network-level storage. Optogenetic silencing of engram cells during retrieval abolishes behavioral expression of the memory, providing causal evidence that specific neuron populations constitute the substrate for long-term memory.^[73] Recent studies as of 2025 have expanded the cellular basis to include non-neuronal elements, with glial cells playing active roles in synaptic plasticity and long-term memory. Astrocytes support LTP through gliotransmitter release (e.g., glutamate, ATP) and regulation of extracellular ion balance, while microglia facilitate synapse refinement via pruning and cytokine signaling, influencing circuit maturation and memory consolidation. For instance, astrocytic mRNA translation controls hippocampal plasticity in fear memory tasks, and microglial interactions bidirectionally tune synaptic remodeling.^[74]^[75]

Retrieval and Forgetting

Retrieval Mechanisms

Retrieval of long-term memories involves accessing stored information through various processes that depend on cues and contextual factors. Two primary types of retrieval are recall and recognition. In recall, individuals actively retrieve information without external aids; free recall requires generating the information entirely from memory, while cued recall provides partial prompts to facilitate access.^[76] Recognition, by contrast, involves identifying previously encountered information from a set of options, often relying on a sense of familiarity rather than detailed reconstruction.^[76] The tip-of-the-tongue (TOT) phenomenon exemplifies challenges in recall, where an item is temporarily inaccessible despite a strong feeling that it is known, accompanied by partial retrieval of related details like the first letter or syllable length.^[77] This state reflects a partial activation of the memory trace, with metacognitive awareness of the impending retrieval.^[77] Context-dependent memory enhances retrieval when environmental cues present during encoding match those at recall, as demonstrated in Godden and Baddeley's 1975 study with scuba divers. Divers who learned word lists either on land or underwater recalled more words when tested in the same environment (mean ≈12.5 words) compared to a different one (mean ≈8.5 words), representing about 40-50% better performance.^[78] This aligns with the encoding specificity principle, where retrieval effectiveness increases when cues overlap with those encoded into the memory. State-dependent memory similarly improves access when internal states, such as mood or physiological conditions, align between encoding and retrieval. For instance, memories formed in a specific mood are more readily retrieved in that same mood, as shown in experiments inducing happiness or sadness.^[79] Drug-induced states, like those from alcohol or amphetamines, also produce state-dependent effects, with recall enhanced when the drug state matches the learning phase.^[80] Neurally, retrieval engages interactive loops between the prefrontal cortex and hippocampus, where the prefrontal cortex initiates search processes and the hippocampus reactivates stored engrams based on cues.^[81] These circuits support controlled retrieval, integrating contextual information to guide access to episodic details.^[82]

Forgetting and Interference

Forgetting in long-term memory refers to the gradual decline in the accessibility or accuracy of stored information over time, distinct from mere retrieval failures. One of the earliest empirical demonstrations of this process came from Hermann Ebbinghaus's self-experiments in 1885, where he memorized lists of nonsense syllables and measured retention at varying intervals. His results revealed the forgetting curve, an exponential decay in recall accuracy that is rapid initially but levels off over longer periods, suggesting that long-term memories stabilize against further rapid loss after initial consolidation.^[83] A primary mechanism underlying this forgetting is interference, where competing memories disrupt the retention or retrieval of target information. In proactive interference, previously learned material hinders the encoding or recall of new information; for instance, in paired-associate learning tasks, prior word pairs (e.g., cat-dog) can cause confusion when learning new associations (e.g., cat-house), reducing accuracy for the newer pairs. Conversely, retroactive interference occurs when subsequent learning impairs access to older memories, as seen in studies where interpolating a second list of paired associates after initial learning leads to poorer recall of the first list, with interference effects persisting into long-term retention tests. These effects, first systematically explored in the mid-20th century, highlight how overlapping similar content in memory traces exacerbates forgetting, particularly when cues are ambiguous.^[84] Beyond passive interference, motivated forgetting involves intentional or unconscious efforts to suppress unwanted memories, serving emotional regulation. Repression, a concept originating in Sigmund Freud's psychoanalytic theory, posits that distressing memories are actively excluded from conscious awareness to protect the psyche, though empirical validation remains debated.^[85] In contrast, modern cognitive models emphasize retrieval suppression, where individuals deliberately inhibit memory activation; neuroimaging studies show this engages prefrontal cortex control to downregulate hippocampal activity, leading to reduced recall of suppressed items even on independent probes. This process, demonstrated in think/no-think paradigms, can weaken memory traces over repeated suppression, contributing to long-term inaccessibility.^[86] Contemporary research views forgetting not merely as a deficit but as an adaptive process that optimizes memory efficiency by eliminating redundant or irrelevant information. Synaptic pruning, a mechanism involving the selective weakening and elimination of unused neural connections, facilitates this by refining long-term memory networks, as evidenced in studies linking it to developmental and adult plasticity for behavioral flexibility.^[87] For example, long-term depression (LTD) during sleep-like states prunes synapses associated with outdated memories, preventing overload and enhancing prioritization of salient information, with 2020s investigations confirming its role in countering interference for adaptive learning.^[88] This perspective reframes forgetting as essential for cognitive economy, aligning with evolutionary pressures to maintain efficient neural resources.^[89]

Impairments and Disorders

Amnesia and Brain Injuries

Amnesia resulting from brain injuries often disrupts the formation and retrieval of long-term memories (LTM), with distinct patterns observed in anterograde and retrograde forms. Anterograde amnesia manifests as an inability to form new LTM following the injury, while retrograde amnesia involves the loss of memories acquired prior to the event. These impairments highlight the vulnerability of neural circuits involved in memory consolidation and storage, particularly those centered around the hippocampus.^[90] A classic example of anterograde amnesia is the case of Henry Molaison (H.M.), who underwent bilateral medial temporal lobe resection in 1953 to treat severe epilepsy, removing much of his hippocampus, amygdala, and parahippocampal gyrus. Post-surgery, H.M. exhibited profound anterograde amnesia, unable to retain new declarative information beyond a few minutes despite intact immediate recall and preserved procedural learning, such as mirror-tracing tasks. This impairment persisted for decades, demonstrating the hippocampus's critical role in transferring short-term memories to LTM.^[35]^[91] Similarly, Clive Wearing, a British musician, developed severe anterograde amnesia after contracting herpes simplex encephalitis in 1985, which caused bilateral damage to his medial temporal lobes. Wearing cannot form new episodic or semantic memories, experiencing a continuous sense of "waking up" every few seconds, though he retains premorbid procedural skills like piano playing and sight-reading short musical passages. His case underscores how focal lesions can selectively abolish LTM encoding while sparing remote, overlearned abilities.^[92] Retrograde amnesia, in contrast, erodes pre-existing LTM, often following Ribot's law, which posits a temporal gradient where recent memories are more susceptible to loss than remote ones due to their less consolidated state. This gradient has been observed in patients with temporal lobe injuries, where memories from years or decades prior remain relatively intact, while those from the preceding 1-2 years are severely disrupted. Evidence from case studies supports this pattern, attributing it to the differential stability of synaptic connections in memory traces. Traumatic brain injury (TBI) frequently induces both anterograde and retrograde amnesia through mechanisms like diffuse axonal injury (DAI), which shears white matter tracts connecting the hippocampus to cortical regions essential for LTM. In moderate to severe TBI, DAI disrupts hippocampal signaling, leading to post-traumatic amnesia (PTA) durations ranging from days to weeks, with longer PTA correlating to poorer LTM outcomes. For instance, spatial memory deficits persist up to a year post-injury in animal models simulating human DAI, reflecting impaired consolidation pathways.^[93] Case studies and neuroimaging provide robust evidence for these impairments. Lesion network mapping of 53 amnesia cases, including strokes and TBIs, reveals a common circuit involving the hippocampal subiculum and retrosplenial cortex, where over 95% of lesions connect, predicting LTM deficits regardless of exact damage location. Functional MRI in TBI survivors shows reduced connectivity in this network during memory tasks, confirming how injuries fragment LTM retrieval even without direct hippocampal destruction.^[90]

Neurodegenerative Diseases

Neurodegenerative diseases represent a group of chronic, progressive disorders that significantly impair long-term memory through protein misfolding and accumulation, leading to neuronal damage in key brain regions. Alzheimer's disease (AD), the most common such condition, is characterized by the formation of extracellular amyloid-β plaques and intracellular tau tangles, which disrupt synaptic function and neuronal connectivity, particularly in the hippocampus—a critical structure for episodic long-term memory formation and retrieval.^[94]^[95] Early in AD, patients experience profound deficits in episodic memory, such as difficulty recalling recent events or personal experiences, while semantic memory remains relatively preserved initially.^[96] Other neurodegenerative diseases also target specific long-term memory subsystems via distinct pathologies. In Parkinson's disease (PD), degeneration of dopaminergic neurons in the basal ganglia leads to impairments in procedural long-term memory, affecting the ability to retain and execute learned motor skills or habits, though explicit declarative memory is less impacted.^[97]^[98] Huntington's disease (HD), caused by CAG repeat expansions in the huntingtin gene, results in striatal atrophy, particularly in the caudate nucleus, which correlates with retrieval deficits in semantic long-term memory, manifesting as reduced access to factual knowledge and word fluency despite intact storage.^[99]^[100] In early stages of these diseases, implicit forms of long-term memory, such as procedural skills, may show relative preservation compared to explicit episodic recall. The progression of long-term memory impairment in these diseases is gradual, beginning with subtle encoding and retrieval failures that escalate to widespread erosion of memory stores over years. Biomarkers such as elevated cerebrospinal fluid (CSF) tau levels, particularly phosphorylated tau, reflect ongoing neurodegeneration and correlate with the rate of memory decline, aiding early diagnosis and monitoring.^[101]^[102] Recent advances in the 2020s include anti-amyloid monoclonal antibody therapies like lecanemab and donanemab, which target amyloid-β and have demonstrated slowing of cognitive decline, including in long-term memory functions, in early-stage AD patients. Lecanemab, targeting soluble amyloid-β protofibrils, showed modest slowing over 18 months in its phase 3 trial, with extension data as of 2025 indicating continued benefits over four years.^[103]^[104]^[105] These interventions reduce amyloid burden and delay episodic memory loss, though they do not reverse established damage.

References

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Physiology, Long Term Memory - StatPearls - NCBI Bookshelf
Long-term memory is consolidated from short-term to long-term memories, primarily in the hippocampus and stored throughout the cortex.
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[PDF] HUMAN MEMORY: A PROPOSED SYSTEM AND ITS CONTROL ...
A class of models for the trace which can explain the tip-of-the-tongue phenomenon are the multiple-copy models suggested by Atkinson and. Shiffrin (1965). In ...
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Human Memory: A Proposed System and its Control Processes
This chapter presents a general theoretical framework of human memory and describes the results of a number of experiments designed to test specific models.
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Long-Term Memory - an overview | ScienceDirect Topics
Long-term memory is defined as the encoding, maintenance, and retrieval of information over extended periods of time, specifically focusing on declarative or ...
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How Long Term Memory Works - Verywell Mind
Feb 13, 2025 · Long-term memory refers to the lasting storage of information in the brain. Learn about the duration, capacity, and types of long-term memory,
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Long-Term Memory - an overview | ScienceDirect Topics
Long-term memory is defined as the more permanent large-capacity memory store that comprises our knowledge base of previously learned information.
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Long-Term Memory In Psychology: Types, Capacity & Duration
Apr 19, 2025 · Long-term memory (LTM) is the final stage of the multi-store memory model proposed by Atkinson-Shiffrin, providing the lasting retention of information and ...
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Long-Term Memory - Cleveland Clinic
Dec 3, 2024 · Long-term memory is a nearly permanent storage space for learned information and experiences. It can hold memories for years.
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Human memory: A proposed system and its control processes.
Atkinson, RC, & Shiffrin, RM (1968). Human memory: A proposed system and its control processes. In KW Spence & JT Spence, The psychology of learning and ...
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[PDF] The Magical Number Seven, Plus or Minus Two - UT Psychology Labs
George A. Miller (1956). Harvard University. First published in Psychological Review, 63, 81-97. My problem is that I have been persecuted by an integer. For ...
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Visual long-term memory has a massive storage capacity for object ...
Here we show that long-term memory is capable of storing a massive number of objects with details from the image.
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[PDF] LOSS OF RECENT MEMORY AFTER BILATERAL HIPPOCAMPAL ...
patient showed marked anterograde and retrograde amnesia. More recently Milnerand Penfield(1955) have described a memoryloss similar in all respects to that ...
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Two storage mechanisms in free recall - ScienceDirect.com
Two experiments were carried out to test the hypothesis that the bimodal serial position curve in free recall is produced by output from two storage mechanisms.
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[PDF] Implications of Short-Term Memory for a General Theory of Memory 1
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Implications of short-term memory for a general theory of memory
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Human memory: theory and data - Internet Archive
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A Power-Law Model of Psychological Memory Strength in Short
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A power-law model of psychological memory strength in short
A classic law of cognition is that forgetting curves are closely approximated by power functions. This law describes relations between different empirical ...Missing: single | Show results with:single
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A distributed memory model for serial-order information
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Serial order effects in short-term memory. - APA PsycNet
STUDIED SERIAL ORDER EFFECTS IN SHORT-TERM MEMORY (STM). THE MOST GENERAL FINDING WAS THE CONSPICUOUS OCCURRENCE OF THE U-SHAPED BOWED SERIAL POSITION CURVE ...
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TODAM2: a model for the storage and retrieval of item, associative ...
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[PDF] Recency-Sensitive Retrieval Processes in Long-Term Free Recall¹
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(PDF) The Demise of Short-Term Memory Revisited: Empirical and ...
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Physiology, Explicit Memory - StatPearls - NCBI Bookshelf - NIH
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[PDF] Episodic and Semantic Memory - Alice Kim, PhD
Kintsch thinks of semantic memory as an organized internal lexicon that represents a person's knowledge of. Page 4. 384/E. TULVING language and that can serve ...
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Autobiographical memory and autonoetic consciousness: triple ...
Based on the first definition of episodic memory (Tulving, 1972, 1983), autobiographical memory has been regarded as episodic in nature. Autobiographical memory ...
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Memory and decision making - PMC - NIH
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8.1 How Memory Functions - Psychology 2e | OpenStax
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Conscious and Unconscious Memory Systems - PMC
Nondeclarative memory (sometimes termed implicit memory) refers to a collection of abilities that are expressed through performance without requiring conscious ...
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Priming and Human Memory Systems | Science
Priming is a nonconscious form of human memory, which is concerned with perceptual identification of words and objects.
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Preserved Learning and Retention of Pattern-Analyzing Skill in ...
Amnesic patients acquired a mirror-reading skill at a rate equivalent to that of matched control subjects and retained it for at least 3 months.
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Conditioning in amnesic patients - ScienceDirect.com
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The Legacy of Patient H.M. for Neuroscience - PMC - NIH
... (Scoville and Milner, 1957). This publication became one of the most cited papers in neuroscience (nearly 2500 citations) and is still cited with high frequency.
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[PDF] A New Look at Habits and the Habit–Goal Interface - USC Dornsife
Additional evidence for the idea that habits are not a form of automatic goal pursuit comes from Neal, Pascoe, and Wood's. (2007) research that manipulated the ...
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Levels of processing: A framework for memory research
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Interactions between attention and memory - ScienceDirect.com
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Mechanisms of emotional arousal and lasting declarative memory
Extensive research in animals implicates stress hormones and the amygdaloid complex as key, interacting modulators of memory consolidation for emotional events.
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Distributed practice in verbal recall tasks: A review and quantitative ...
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Mnemonic Devices: Classification, Characteristics, and Criteria
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The magical number seven, plus or minus two: Some limits on our ...
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Benefits of multisensory learning - PubMed
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Review Sleep—A brain-state serving systems memory consolidation
Apr 5, 2023 · Although long-term memory consolidation is supported by sleep, it is unclear how it differs from that during wakefulness.
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Temporal Flexibility of Systems Consolidation and the Synaptic ...
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Slow-wave sleep and the consolidation of long-term memory
Slow-wave sleep (SWS) has been shown to play an important role in the reinforcement of declarative memory. A dialogue between the neocortex and hippocampus ...
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Low acetylcholine during slow-wave sleep is critical for declarative ...
During slow-wave sleep (SWS), however, declarative memory consolidation is particularly strong, while acetylcholine levels in the hippocampus drop to a minimum.
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The REM Sleep–Memory Consolidation Hypothesis - PMC
It has been hypothesized that REM (rapid eye movement) sleep has an important role in memory consolidation. The evidence for this hypothesis is reviewed and ...
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The effect of selective REM-sleep deprivation on the consolidation ...
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Reactivation of hippocampal ensemble memories during sleep
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Reactivation of Hippocampal Ensemble Memories During Sleep
Simultaneous recordings were made from large ensembles of hippocampal "place cells" in three rats during spatial behavioral tasks and in slow-wave sleep ...
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The Common Effects of Sleep Deprivation on Human Long-Term ...
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The Devastating Effects of Sleep Deprivation on Memory
May 15, 2023 · According to a study, prolonged sleep disruption inhibits the regeneration of hippocampal neurons and impairs memory. Moreover, sleep disruption ...
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An update on recent advances in targeted memory reactivation ...
Apr 15, 2024 · TMR is a noninvasive tool to manipulate memory consolidation during sleep. TMR builds on the brain's natural processes of memory reactivation during sleep.
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Disarming emotional memories using targeted memory reactivation ...
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The Episodic Memory System: Neurocircuitry and Disorders - Nature
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Interplay of hippocampus and prefrontal cortex in memory - PMC
the hippocampus and the prefrontal cortex — that support ...
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Where are memories stored in the brain? - Queensland Brain Institute
There are three areas of the brain involved in explicit memory: the hippocampus, the neo-cortex and the amygdala. Hippocampus. The hippocampus, located in the ...
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Retire Papez circuit: space, memory, attention
After more than 80 years, Papez serial circuit remains a hugely influential concept, initially for emotion, but in more recent decades, for memory.
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An Update to the Original Papez Circuit of the Human Limbic System
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The role of the basal ganglia in learning and memory
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Human brain activity and functional connectivity associated with ...
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Hemispheric lateralization of verbal and spatial working memory ...
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NMDA receptors - their role in long-term potentiation - ScienceDirect
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Arc in synaptic plasticity: from gene to behavior - PubMed Central
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DNA methylation-mediated control of learning and memory
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DNA methylation: a permissive mark in memory formation and ...
This review highlights the requirements for the enzymes responsible for DNA methylation and demethylation in memory encoding. In addition, it describes the ...
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Identification and optogenetic manipulation of memory engrams in ...
Jan 17, 2014 · We will review recent advances in memory engram research that combine transgenic and optogenetic approaches to reveal the underlying neuronal substrates.
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The neurobiological foundation of memory retrieval - PMC
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Tip-of-the-tongue (TOT) states: retrieval, behavior, and experience
Jan 4, 2011 · The tip-of-the-tongue state (TOT) is the feeling that accompanies temporary inaccessibility of an item that a person is trying to retrieve.
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CONTEXT‐DEPENDENT MEMORY IN TWO NATURAL ...
In a free recall experiment, divers learnt lists of words in two natural environments: on dry land and underwater, and recalled the words in either the ...
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Mood state-Dependent Retrieval: The Effects of Induced Mood on ...
Five experiments are reported investigating the effects of mood on learning and recall. Mood-state-dependent retrieval was observed in Experiment 1a (using ...
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State-Dependent Memory - an overview | ScienceDirect Topics
The drugs that most reliably produce state-dependent retrieval effects (such as alcohol and amphetamines) are accompanied by large mood changes. This led Bower ...
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Prefrontal-Hippocampal Interactions in Memory and Emotion - NIH
Dec 15, 2015 · The hippocampal formation (HPC) and medial prefrontal cortex (mPFC) have well-established roles in memory encoding and retrieval.
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Multiple modes of hippocampal-prefrontal interactions in memory ...
The hippocampus and prefrontal cortex are critical for learning and memory-guided behavior, but neural mechanisms underlying their coordinated operation are ...
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Ebbinghaus (1885/1913) Chapter 1
Hermann Ebbinghaus (1885). Translated by Henry A. Ruger & Clara E. Bussenius (1913) Originally published in New York by Teachers College, Columbia University.
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[PDF] Interference theory: History and current status - University of Waterloo
This chapter surveys the history of research on interference in learning and memory from the earliest empirical work at the dawn of experimental psychology to ...
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Repression in Psychology - Verywell Mind
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https://www.sciencedirect.com/topics/neuroscience/state-dependent-memory
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Pruning recurrent neural networks replicates adolescent changes in ...
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The times they are a-changin': a proposal on how brain flexibility ...
Dec 27, 2022 · Here, we will focus on studies showing that downward synaptic plasticity is required for adulthood memory formation as well as for post- ...
[83]
Remembering to Forget: A Dual Role for Sleep Oscillations in ...
The same sleep rhythms that consolidate new memories, in the cortex and hippocampus, simultaneously organize the adaptive forgetting of older memories in these ...
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A human memory circuit derived from brain lesions causing amnesia
Aug 2, 2019 · We derive a human memory circuit using 53 case reports of strokes causing amnesia and a map of the human connectome (n = 1000).
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LOSS OF RECENT MEMORY AFTER BILATERAL HIPPOCAMPAL ...
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Musical memory in a patient with severe anterograde amnesia - NIH
Perhaps the most well-known of these cases is Clive Wearing, a renowned ... Piano playing in Alzheimer's disease: Longitudinal study of a single case.
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Cognitive Impairment following Traumatic Brain Injury - NCBI - NIH
Both retrograde and anterograde memory deficits are common following TBI. Memory function seems to be particularly vulnerable to head injury, and Levin and ...
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What Happens to the Brain in Alzheimer's Disease?
Jan 19, 2024 · It appears that abnormal tau accumulates in specific brain regions involved in memory. Beta-amyloid clumps into plaques between neurons. As the ...
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Memory Performance is Related to Amyloid and Tau Pathology in ...
This study implicates both amyloid deposition and tau pathology in the hippocampus as an early and late cause of decline in memory function over time in AD.
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Hippocampus and its involvement in Alzheimer's disease: a review
Feb 1, 2022 · The hippocampus is one of the brain areas affected by Alzheimer's (AD). In the early stages of AD, the hippocampus shows rapid loss of its tissue.
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Retrograde Procedural Memory in Parkinson's Disease
Feb 9, 2022 · We hypothesized that retrograde procedural memory is impaired in people with Parkinson's disease compared to an age- and gender-matched control group.
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Motor Learning Deficits in Parkinson's Disease (PD) and Their Effect ...
Studies have shown that procedural and spatial memory is especially affected in PD, while verbal and episodic memories remain largely intact (103, 104, 106, 107) ...
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The disparate effects of Alzheimer's disease and Huntington's ...
According to one view, AD disrupts the storage of semantic memories, whereas HD disrupts the retrieval of semantic memories. Dissenters argue that AD, like HD, ...
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Semantic, phonologic, and verb fluency in Huntington's disease - PMC
Thus, various studies have indicated poor performances on phonologic and semantic fluency tests in individuals with Huntington's disease (HD).
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Tau as a biomarker of neurodegenerative diseases - PMC
As peripheral biomarker of Alzheimer's disease in the cerebrospinal fluid, Tau proteins are now validated for diagnosis and predictive purposes. For the future, ...
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CSF Biomarkers for Alzheimer's Disease Diagnosis - PMC
Thus far, three CSF biomarkers, Aβ42, total-tau (t-tau), and phosphorylated-tau (p-tau), have been found to have the highest diagnostic potential. Biomarkers ...
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Lecanemab in Early Alzheimer's Disease
Nov 29, 2022 · Lecanemab reduced markers of amyloid in early Alzheimer's disease and resulted in moderately less decline on measures of cognition and function than placebo at ...
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Re-evaluation of the efficacy and safety of anti-Aβ monoclonal ...
Apr 16, 2025 · Lecanemab/Donanemab can improve memory, cognitive function, and daily living abilities in patients with early AD, significantly reduce the composite score of ...