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3. Memory Rudolf Cardinal [email protected] University Lecturer in Clinical Informatics, Department of Psychiatry, University of Cambridge Honorary consultant psychiatrist, Liaison Psychiatry Service, CPFT & CUH Cambridge MRCPsych course Tue 23 Oct 2018, pm MRCPsych syllabus (2018) includes: Overview ● Memory is the ability to retain information. Very broad term. Perceptual, motor. ● Types of memory and some other key aspects of the psychology of memory. ● Some neurobiology (not comprehensive, e.g. cerebellar memory). ● Some exam-style questions. Types of memory old terms: primary memory = STM secondary memory = LTM Morris (2001), after Tulving The ‘sensory store’ — e.g. visual ‘iconic’ memory P H O G M Y V Q X L C U ● Brief (50ms) visual presentation → mask → delay → signal to recall an unpredictable subset. Access to 9–10 letters. ● A large amount of information is available for a very short time (about 0.5s). After this: STM, and access to 4 letters. ● Auditory version (“echoic” store): about 2 s. Sperling (1960) STM is very S … and of limited capacity Miller’s magical number 7 ± 2, digit span, and chunking: 8164912 snailcatdogfishthrushbatworm mstoarfhbcadagstnriwtihoulsh Miller (1956); Peterson & Peterson (1959) Baddeley & Hitch (1974) model of working memory ● The visuospatial sketchpad and the phonological loop are “slave systems”. Primacy and recency (serial position) effects primacy effect recency effect “40-1” = 40 items for 1 s each Theories include: ● Dual-store models: e.g. primacy relates to LTM storage; recency relates to STM. ● Single-store models: e.g. relating to relative strength of memory trace; differential rehearsal effects; etc. Murdock (1962); Sederberg et al. (2008) Forgetting and remembering; terms Forgetting: not just decay, but ● decay [with time; STM] interference by new material? ● interference [by other things] An example (but not a simple ● displacement [from STM] one)… ● motivated forgetting [active, even if unconscious, like ‘repression’] ● remembering [vague] ● search / retrieval [from LTM] ● recognition [something is familiar/known] ● recall [free, cued] = retrieval + recognition of recalled item? ● retrieval failure / tip-of-the-tongue state ● cue-dependent forgetting / state- dependent memory ● rehearsal ● consolidation [STM to LTM] / reconsolidation [advanced] e.g. Jenkins & Dallenbach (1924) State-dependent memory Does the context become part of the memory? Recall is easier if you’re in the same state or context as that in which you learned. Referred to as state- dependency. Other catchphrase: “cue-dependent forgetting”. Examples: SS same state • room 1 versus room 2 DS different state E encoding • on land versus underwater R retrieval • sober versus drunk (for tests of explicit memory) e.g. Abernathy (1940) rooms; Godden & Baddeley (1975) underwater; Duka et al. (2001) drunk Schemata (schemas) and memory distortion ● memory as active reconstruction ● incorporation of existing knowledge ● the use of prior knowledge is vital (even for perception) ● but can produce distortion to fit existing schema e.g. ● Bartlett (1932): Native American “War of the Ghosts” to UK students, with serial reproduction by each individual. ● Allport & Postman (1947): the passing of stories down a chain of people. ● Treadway & McCloskey (1987): the corruption of the Allport & Postman study by professional psychologists, giving legal testimony relating to eyewitness accounts, who hadn’t read the original. Episodic versus semantic memory Semantic memory… categories Semantic memory: cortical, distributed, related to perception? Priming Meyer & Schvaneveldt (1971) Procedural versus declarative memory after Dickinson (1980) Human amnesia from Markowitsch (1995) H.M.’s bilateral medial temporal lobe resection on MRI H.M. normal brain EC entorhinal cortex, MMN medial mammillary nucleus; A amygdala; H hippocampus CS collateral sulcus; PR perirhinal cortex 1953 operation: Scoville & Milner (1957) J Neurol Neurosurg Psych 20: 11 MRI: Corkin et al. (1997) J Neuro 17: 3694 Preserved abilities in medial temporal lobe amnesia Profound anterograde amnesia. Impaired recognition. Some retrograde amnesia (temporally graded). But • IQ normal • Could learn mirror-writing (Milner 1962, 1965) and similar motor skills day-by-day, despite inability to remember that he’d done it before. • Learned a perceptual learning task (recognition of words from incomplete fragments) • Improved with practice on the Tower of Hanoi task (Cohen 1984) • Short-term memory: normal digit span and visual immediate memory • Priming normal (typical of amnesiacs, see Aggleton & Brown 1999) McCarthy & Warrington (1990) The medial temporal lobe: hippocampus, amygdala, fornix The hippocampal formation in cross-section Martin (1989, p391) The hippocampal formation in cross-section (approx.!) perforant path (EC DG) (includes entorhinal cortex) Martin (1989, p391, modified) The contribution of medial temporal lobe structures ● The delayed non-matching to sample task: which of two objects is new? ● Aspiration of medial temporal lobe structures (esp. hippocampus) impairs DNMTS. So you might think e.g. the hippo. is required. ● But excitotoxic lesions of the hippocampus don’t; the rhinal cortex (includes entorhinal, perirhinal cortex) is critical instead. ● Perirhinal cortex is the first polymodal ventral stream area. It may encode feature conjunctions. ● The hippocampus is important for navigation (place cells; 2014 Nobel Prize [O’Keefe/Moser/Moser]; the hippocampus as a cognitive map; Morris water maze; taxi drivers). ● Also for encoding scenes (monkeys, Raiders of the Lost Ark). ● Common role may be relational coding (e.g. what/where/when conjunction required for episodic memory). Squire & Zola-Morgan (1991); Murray & Mishkin (1998); Murray & Bussey (1999); Bussey & Saksida (2002); O’Keefe & Nadel (1978); Morris et al. (1982); Maguire et al. (1997); Gaffan (1992); Eichenbaum et al. (1999) Long-term potentiation (LTP) as a cellular model of memory ● Glutamate receptor types include AMPA, NMDA ● NMDA receptors are voltage- and ligand-gated ● With sufficient depolarization [postsynaptic activity] plus glutamate [presynaptic activity], they will open and let calcium in, leading to synaptic plasticity. ● Simple mechanism for Hebb’s postulate (cells that fire together, wire together). ● Discovered in the hippocampus (Bliss & Lømo, 1973). Patient N.A.: fencing foil (up nostril) to diencephalon (Normal brain! Approximate area of damage in N.A. circled.) Diencephalon: thalamus, hypothalamus, epithalamus The Delay–Brion (= Papez) circuit: hippocampus fornix mammillary bodies mammillothalamic tract anterior thalamus (thalamus) (posterior hypothalamus; doesn’t connect much with other hypothalamic regions) N.B. Wernicke–Korsakoff encephalopathy (myelin stain; from Martin, 1991; also Aggleton, 2014) Semantic dementia: impaired semantic, preserved episodic? 1 semantic task — name a familiar object ‘episodic’ task — recognize an object (‘perceptually identical’) mixed task — recognize a different example of an object (‘perceptually different’) Graham et al. (2000) Semantic dementia: impaired semantic, preserved episodic? 2 a subtype of frontotemporal dementia – the impaired semantic performance temporal end normal object recognition; failure to recognize a different example of the same kind of object Graham et al. (2000) Semantic dementia: damage to a simple associative net? computational model patient with progressive semantic dementia Moss et al. (2002) Moving memories. Sleep. ● Graded retrograde amnesia following medial temporal lobe lesions. ● Gradual transfer of memories from hippocampus/MTL to elsewhere? ● Hippocampus ‘replays’ pattern to cortex, allowing cortical strengthening? ● Role of sleep. ● Hippocampus ‘replays’ activity during sleep. ● Sleep aids consolidation of procedural memory (e.g. finger- tapping sequence). ● Sleep aids acquisition of insight into hidden task rules. Scoville & Milner (1957); Squire et al. (2001); Louie & Wilson (2001); Fischer et al. (2002); Wagner et al. (2004) Consolidation and reconsolidation consolidation ● potential use for ‘erasure’ of pathological reconsolidation memories ● old study of conscious ECT for OCD ● consideration of drug/ cue-exposure combinations Nader (2003) Habit learning A double dissociation: Parkinson’s disease and amnesia (1) • Task 1 (probabilistic classification): one to three cards are shown. The subject must predict sunshine or rain. Feedback is provided (correct/incorrect). One cue is associated with sunshine on 25% of occasions; one on 43% of occasions; one 57%; one 75%. • Task 2 (declarative): memory for features of the game (screen layout, cues, etc.) is tested with four-way multiple-choice questions. Knowlton et al. (1996) A double dissociation: Parkinson’s disease and amnesia (2) • PD patients: impaired on probabilistic classification task, not declarative. (PD* = severe.) • Amnesic patients (with bilateral hippocampal damage or midline diencephalic damage): impaired on declarative task, not probabilistic classification. Knowlton et al. (1996) Memory and the prefrontal cortex Delayed response task: PFC active during the delays Friedman & Goldman-Rakic (1988); task by Hunter (1913); Fuster & Alexander (1971) Working memory: PFC maintains posterior cortex activity? Two-colour DMTS Coloured areas: activity of IT neurons. Black bars: behavioural performance. Four-colour DMTS Fuster & Alexander (1970); Fuster (1995) Best of five: Which reflects semantic memory? A. remembering where you bought your bicycle