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Memory – Outline • Long-term memory (LTM) – Types: episodic, semantic – Stages • Memory loss (amnesia) – Brain regions & diseases • Implicit Long-term memory • Spatial memory Possible architecture of: Long-term Memory Implicit Memory Explicit Memory Motor Skills Semantic Memory Episodic Memory ? Affected by Amnesia Perceptual learning Other types Classical Conditioning Episodic Memory = Relational Memory • • • • What did you do at home last Wednesday? A temporal context (last Wednesday) A physical context (home) A set of memory events (turkey, annoying parents, tiredness, watching the game) A relation between the events and the context Stages of Explicit Long-Term Memory Encoding Acquire information via perceptual system Storage Establish a ‘permanent record’ Retrieval Reactivate the permanent record Object Representation, based on activation of perceptual & motor areas Stimulus banana Perceptual Processes Episodic Memory Encoding word action shape In Context: ‘today’ ‘at home’ location temporal lobe is the hub that color relates all object’s properties & context into an integrated A medio-temporal lobe lesion causes Amnesia! memory Storage of Memories (consolidation) Episodic Memory Encoding Storage (Consolidation) Old memory (10 years later) (new memory) In Context: ‘today’ ‘at home’ temporal lobe is the hub that relates all object’s properties into an integrated memory temporal lobe is not needed anymore Consolidation hypothesis of amnesia Episodic Memory Encoding Storage (Consolidation) Old memory (10 years later) (new memory) In Context: ‘today’ ‘at home’ RECENT MEMORIES ARE LOST temporal lobe is the hub that relates all object’s properties into an integrated memory OLD MEMORIES ARE SPARED temporal lobe is not needed anymore Retrograde Amnesia (Pre-Damage) Old memory (spared) Anterograde Amnesia (Post-Damage) Recent memory (lost) 1960’s 1970’s New memories (not encoded) 1980’s Brain Damage (1975) 1990’s Semantic memory: a deficit in amnesia? • Semantic memory is usually spared in amnesia • but most is acquired early in life (‘what’s an apple?’) • May be it is spared because its an old memory • Test: would amnesics learn new knowledge? – ipod – Britney spears? ‘Elvis’ ? ??? ‘Chaplin’ 1930’s 1950’s 1960’s 1980’s Brain Damage (1970) ??? 1990’s Anatomy of episodic memory Lateral View temporal lobe is the hub that relates all object’s properties into an integrated concept Ventral View Coronal View Mediotemporal lobe, area removed in amnesic patient H.M. Anatomy of episodic memory: Neuroimaging evidence Task: Read these words and try to remember them Task: Recall as many words as possible Porcupine Lawyer Peace Simple Octagon Walk .. .. Trigger Applause Porcupine ------Peace ----Octagon Walk .. .. -------Applause 30’ delay ENCODING Method: use fMRI to measure brain activity RETRIEVAL Data analysis: Contrast brain activity during encoding of: words that would later be recalled (peace) vs. words that would be forgotten (simple) Results: activation of hippocampus while encoding to-be recalled words (peace) Anatomy of episodic memory: Neuroimaging evidence % of participants that correctly recallled the word hippocampus activity Conclusion: fMRI in normal subjects provide convergent evidence for the role of hippocampus in episodic memory encoding Results: activation of hippocampus while encoding to-be recalled words (peace) Anatomy of episodic memory: evidence from infantile amnesia (?) • We are unable to recall personal experiences that took place before about 3 years of age, • probably because the hippocampus is not yet developed, so we can’t retain new explicit memories Neuroanatomy of episodic memory: Evidence from a famous case • Patient H.M. – He suffered from seizures, which were treated in 1953, at the age of 29, with – Bilateral removal of medio-temporal lobe – Surgery led to anterograde amnesia – He was spared in other domains • digit span, • conversation, • motor learning – He died in 2008. Neuroanatomy of episodic memory: Evidence from a famous case • Patient H.M.: A detailed comparison of HM’s brain to that of normal brain Causes of Amnesia Fornix Mammillary body (Korsakoff Patients) Hippocampal formation – HM, Alzheimer’s disease Coronal View Causes of Amnesia Atrophy of mammillary bodies due to vitamin depletion (Korsakoff’s syndrome) Usually in patients with chronic alcoholism Mammillary body Mammillary body Causes of amnesia Coronal View Saggittal Cut -Lack of Oxygen (Anoxia) usually due to cardiac arrest Enlarged view: Normal hippocampus Anoxia Causes of amnesia Alzheimer’s disease (midtemporal lobe atrophy) Coronal view Lateral view Dorsal view Causes of Amnesia: - Alzheimer’s Disease • Gradual decline (dementia) • It affects: – 13% of > 65 y-old, – 42% of > 85 y-old • Brain pathology – Plaques and tangles • early = hippocampus, • later = cortex and others – Loss of acetylcholine cells Possible architecture of: Long-term Memory Implicit Memory Explicit Memory Motor Skills Semantic Memory Episodic Memory Affected by Amnesia Perceptual learning Other types Classical Conditioning Implicit memory: Perceptual priming Implicit memory: motor sequence learning task Implicit Memory: Mirror drawing task Implicit memory: classical conditioning – Eyeblink response implicit memory: rule learning • Tower of Hanoi game – Amnesics cannot remember playing, but they get faster at solving the game. Start Position Goal 2 (2 moves) Goal 10 (5 moves) Implicit memory: Conceptual Priming • Study phase: – “is this word pleasant or not? “rape”, “love”, “sincere”, …..” • Test phase: A. Free Recall: Name the words on the list B. Word Stem Completion: Complete “sco_ _ _” • Results: A. Impaired free recall, but B. Normal stem completion: - as likely as normals to complete stems with studied words (Graf, Squire, & Mandler, 1984) Summary • Medial Temporal Lobe Amnesics – – • Explicit memory: unable to create Implicit memory: relatively intact So far: single dissociation – 2 possibilities: 1. Implicit/Explicit 2 independent systems 2. Implicit simply easier, relies on residual processing of a single, partially damaged system – Double dissociation would support claim of 2 independent systems Evidence for double dissociation: Patient MS – Intractable epilepsy: – surgery removed right occipital lobe (BA 17,18, part of 19). – Hemianopic (blind in left field) Evidence for double dissociation: Explicit Memory: Recognition Task – Read 24 words --> delay --> read 48 words • 24 from study phase, • 24 new foils – Task: have you seen this word? Results: – Middle Temporal Lobe Amnesics (e.g. HM): impaired – Visual Cortex lesion (patient MS): normal Gabrieli et al. (1995) Evidence for double dissociation: Implicit Memory: Word-stem completion task – Read or Listen 24 words (‘colt’) --> delay --> Read 48 word-stems (‘co__’) • 24 from study phase, (e.g., ‘co___’) • 24 new unrelated stems (e.g., ‘li__”) – Task: complete word stem Results: – Healthy adults: Visual Priming (faster reading of previously read words than previously heard words). – Middle Temporal Lobe Amnesics (HM): Visual Priming – Visual Cortex Lesion (MS): no Visual Priming – Conceptual priming (across-domains) is spared in all groups Gabrieli et al. (1995) Evidence for double dissociation: • Summary – Visual cortex lesion (patient MS): • Spared Explicit memory • Impaired visual priming – Middle temporal lobe lesion (patient HM, Alzheimer) • Impaired Explicit memory • Spared visual priming – Double dissociation: • Two independent systems – one for visual priming (implicit memory), – the other for explicit memory Spatial memory • Anatomy: – Medial temporal lobe: • Hippocampus • Posterior parahippocampal area • Evidence – Human Neuropsychology • HM • London taxi drivers – Animal research Medial temporal lobe (MRI) Folded shape stained hippocampus 3. Posterior 4. Amygdala 1. Hippocampus 2. Perirhinal cortex parahippocampal cortex Mastering London Topography Changes Hippocampal Structure in Taxi Drivers (humans) Bird species with good spatial memory have larger hippocampi Spatial memory: Animal research Spatial memory: Animal research Place fields are stationary over time Place cell: A neuron that becomes active when the animal is in a particular location in the environment; most typically found in the hippocampal formation (in dorsal hippoc, which is analogous to human post hippoc) Notes • Episodic Memory: specific personal experiences from a particular time and place. For example, I remember that “Ted Kennedy died (last week)”. ‘porcupine was on the list I read 20 mins ago’ • Semantic memory: world knoweldge. The sky is blue. Dogs are mammals, are pets, have four legs, • Perceptual learning: getting accustomed to Diego’s accent • Classical conditioning: pavlov’s dog Spared slides Recall vs. Recognition • Recognition relies on – Recollection: remembering specific info from the study episode (i.e., is context dependent – Familiarity: context free – Recognition Amnesia Other causes of amnesia include - Anoxia (CA1 atrophy) Interestingly, a genetic mutation inactivating NMDA receptors in CA1 leads to: -Lack of Long-term potentiation -Poor spatial memory in the watermaze -Large & unfocused spatial receptive fields Some strategies for communicating with someone who has Alzheimer’s Disease Problem Strategy Asking the same questions over & over Distract with other subjects, avoid arguing Denying memory problems, blaming others Don’t force recognition of memory problems; be sympathetic Failing to recognize familiar people, objects Agree things look different, calmly remind Memory Retrieval Sensory Memory Short-term Working Memory Long-term Memory (knowledge) Encoding 200 ms ‘Vision’ 20 secs Implicit memory Working Memory fMRI study of WM, showing distinct activation to faces and scenes Working Memory & Long-term memory in a single task: the Free Recall Task (& serial position function) Subjects: - hear items (usually 10-40 words), then - they say or write all the items they can remember, in any order. Serial Position Function Probability of reporting the item ? 12 ……… Position in Original List 30 1. Monster 2. Camera 3. Tricycle 4. Melon 5. Window 6. Guest 7. Quiet 8. Cherish 9. Waiting 10. Rabbitt 11. Computer 12. Child 13. Chicken 14. Ghost 15. Slave Faster presentation rate 30 secs filled delay Patient HM Primacy Recency Privileged rehearsal better LTM encoding STM contribution Relational Memory • • • • What did you do at home last Wednesday? A temporal context (last Wednesday) A physical context (home) A set of memory events (turkey, annoying parents, tiredness, watching the game) A relation between the events and the context (relational memory) SPARE SLIDES From Patterson, 2007, NatNeuroRev 60 Copyright © Allyn & Bacon 2007 61 Copyright © Allyn & Bacon 2007 62 Copyright © Allyn & Bacon 2007 Delayed non-match to sample • Familiarity vs. explicit memory Long-term Memory Systems Implicit Memory Explicit Memory Motor Skill Learning Priming Semantic Memory Episodic Memory Classical Conditioning Instrumental Learning Chapter 13: Single cell recording Instrumental learning Classical Conditioning (associative Learning) What is the bases of such learning? Synaptic plasticity: “cells that fire together, wire together” (Donald Hebb) Long-term potentiation: Following increased input there is a long-term increase in the excitability of the neuron that receives such input Long-term potentiation in the rat hippocampus