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Neuroscience: Exploring the Brain, 4e. Chapter 24: Memory Systems. Introduction. Learning and memory: lifelong brain adaptation to environment Several similarities between experience-dependent brain development and learning
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Neuroscience: Exploring the Brain, 4e Chapter 24: Memory Systems ..
Introduction • Learning and memory: lifelong brain adaptation to environment • Several similarities between experience-dependent brain development and learning • Similar mechanisms at different times and in different cortical areas • Memories range from stated facts to ingrained motor patterns. • Anatomy: several memory systems • Evident from effects of brain lesions
Types of Memory and Amnesia • Learning: acquisition of new information • Memory: retention of learned information • Declarative memory (explicit) • Facts and events • Nondeclarative memory (implicit) • Procedural memory—motor skills, habits
Types of Procedural Memory • A type of nondeclarative memory • Involves learning a motor response (procedure) • In reaction to sensory input • Occurs in two categories of learning • Nonassociative learning • Associative learning
Types of Nonassociative Learning (a) Habituation (b) Sensitization
Associative Learning • Behavior altered by formation of associations between events • In contrast to changed response to a single stimulus • Classical conditioning (Pavlov) • Pairing of unconditional stimulus with conditional stimulus • Instrumental conditioning (Thorndike) • Associate a response with a meaningful stimulus
Types of Declarative Memory • Working memory • Temporary storage, lasting seconds • Short-term memories—vulnerable to disruption • Facts and events stored in short-term memory • Subset are converted to long-term memories. • Long-term memories • Recalled months or years later • Memory consolidation: process of converting short- to long-term memories
Amnesia • Amnesia: serious loss of memory and/or ability to learn • Causes: concussion, chronic alcoholism, encephalitis, brain tumor, stroke • Limited amnesia (common)—caused by trauma • Dissociated amnesia: no other cognitive deficits (rare) • Retrograde amnesia: memory loss for things prior to brain trauma • Anterograde amnesia: inability to form new memories after brain trauma
Amnesia—(cont.) • Transient global amnesia • Sudden onset of anterograde amnesia • Also inability to recall the recent past • Lasts a shorter period, from temporary ischemia (e.g., severe blow to head) • Symptoms: disoriented, ask same questions repeatedly; attacks subside in couple of hours; permanent memory gap
Working Memory • We pay attention to small fraction of sensory information. • Some sensory information held briefly in working memory. • Small capacity—limited resource • Mostly discarded, some may be converted to long-term memory • A capability of neocortex found in numerous brain locations
Prefrontal Cortex and Working Memory • Primates have a large frontal lobe. • Functions of prefrontal cortex: self-awareness, capacity for planning and problem solving
Wisconsin Card-Sorting Test • To demonstrate problems associated with prefrontal cortical damage
Working Memory Activity in Monkey Prefrontal Cortex Prefrontal cortex
Imaging Working Memory in the Human Brain • Numerous brain areas in prefrontal cortex involved in working memory. • PET study: Six frontal lobe areas show sustained activity correlated with working memory. • Identity task • Location task • Unknown whether working memory for other types of information is held in same or different brain areas
Human Brain Activity in Two Working Memory Tasks Blue; identity Green; both Red; location
Area LIP and Working Memory • Cortical areas outside frontal lobe also involved in working memory. • Lateral intraparietal cortex (area LIP) • Involved in guiding eye movements • Stimulation causes saccades. • Demonstrated in delayed-saccade task in monkeys • Other modality—specific areas of parietal and temporal cortex have analogous working memory responses
The Neocortex and Declarative Memory • Lashley’s rat experiments • Cortical lesions produce memory deficits. • Speculated all cortical areas contribute equally (equipotential) • Equipotential capacity later disproved • But memory engrams can be widely distributed in the brain
Hebb and the Cell Assembly • External events are represented in cortical cells. • Cells reciprocally interconnected reverberation • Simultaneously active neurons—cell assembly • Consolidation by “growth process” • “Fire together, wire together” • Hebb on the engram • Widely distributed among linked cells in the assembly • Could involve neurons involved in sensation and perception
The Medial Temporal Lobes • Important for consolidation and storage of declarative memories • Demonstrated by: • Electrical stimulation in the temporal lobe • Neural recordings from the temporal lobe
Electrical Stimulation of the Human Temporal Lobes • Temporal lobe stimulation • Effects different from stimulation of other areas of neocortex • Penfield’s experiments • Stimulation sensations like hallucinations or recalling past experiences • Temporal lobe: apparent role in memory storage • Caveat: complex sensations reported by minority of patients, all with abnormal brains (epilepsy)
Human Neural Recordings from the Medial Temporal Lobe • Neurons found that preferentially respond to categories • Faces, household objects, outdoor scenes • Invariant neurons—respond to variety of images are structurally or conceptually related • Individual neurons respond selectively to one person’s face. • Many questions remain.
A patient’s hippocampal neuron selectively responds to actress Halle Berry
Temporal Lobectomy and Amnesia (H.M.) • Removal of temporal lobes had no effect on perception, intelligence, personality. • Anterograde amnesia so profound he could not perform basic human activities (and partial retrograde amnesia) • He could not recognize the doctor who studied him for nearly 50 years. • Impaired declarative memory, but spared procedural memory (mirror drawing) • Normal working memory, morality, IQ • Poor imagination
An Animal Model of Human Amnesia • Studies of macaque medial temporal lobe using experimental ablation • Delayed match-to-sample and delayed non-match to sample (DNMS) tests • Recognition memory tasks • Amygdala and hippocampus not significantly involved in recognition memory • Much still unknown about specific brain areas • Collectively, medial temporal structures critical for consolidation of memory
Delayed Non-Match to Sample (DNMS) Task • Medial temporal lobe structures shown important for memory consolidation
The Diencephalon and Memory Processing: The Case of N.A. • Radar technician accidentally stabbed through left dorsomedial thalamus • Less severe amnesia, but like H.M.: anterograde and some retrograde amnesia • Korsakoff’s syndrome: chronic alcoholism—thiamin deficiency • Symptoms: confusion, confabulations, severe memory impairment, apathy • Can lead to lesions in dorsomedial thalamus and mammillary bodies • Suggests mechanisms involved in consolidation distinct from processes that recall memories
Memory Functions of the Hippocampal System • Memory formation, retention, retrieval involve system of interconnected brain areas • Hippocampus involved in various memory functions • Binds sensory information for memory consolidation • Supports spatial memory of location of objects of behavioral importance • Involved in storage of memories for some length of time
Effects of Hippocampal Lesions in Rats (a) Normal rats go down each maze arm for food only once - but not with hippocampal lesions (b) Normal and lesioned rats learn which arms are baited and avoid the rest
Spatial Memory and Place Cells • Learning Morris water maze requires hippocampus. • Place cells fire when animal is in a specific place. • Place fields dynamic
Place Cells in Humans • PET imaging in human brain related to spatial navigation of a virtual town
Grid Cells • Identified in rodent neural recordings • Inthe entorhinal cortex • Unlike place cells • Respond when animal is at multiple locations that form hexagonal grid • Likely also grid cells in human entorhinal cortex • Place cells, grid cells, and hippocampal neurons showing sensitivity for head direction brain region highly specialized for spatial navigation
Hippocampal Functions Beyond Spatial Memory • O’Keefe and Nadel: hippocampus specialized for creating spatial map of environment • Apparent important role in spatial memory • Other hippocampal function theories • Important for working memory • Integrates or associates sensory input • Odor discrimination • Hippocampus links different experiences together.
Consolidating Memories and Retaining Engrams • Declarative memory formation involves system of interconnected brain structures: • Take in sensory information • Make associations between related information • Consolidate learned information • Store engrams for later recall • Components include hippocampus, cortical areas around hippocampus, diencephalon, neocortex, and more.
Two Models of Memory Consolidation • Standard model of memory consolidation • Information from neocortex areas associated with sensory systems sent to medial temporal lobe for processing • Synaptic consolidation, systems consolidation • Multiple trace model of consolidation • Engrams involve neocortex, but even old memories also involve hippocampus. • Multiple memory traces
Reconsolidation • Rat experiments • Reactivating a memory makes it sensitive to change as when first formed (before consolidation) • Reconsolidation: the reactivation effect • Human reconsolidation experiments • Recalling a memory makes it susceptible to change • Hippocampal activity • Profound implications for treatment of stress associated with unpleasant memories