Cbu Human Memory

8/4/2019 Cbu Human Memory

1/35

THE SYSTEMS NEUROSCIENCETHE SYSTEMS NEUROSCIENCE

OF HUMAN MEMORYOF HUMAN MEMORY

Rik HensonRik Henson


2/35

OverviewOverview

A taxonomy of memoryA taxonomy of memory

For each type of memory:For each type of memory:

Definition and Common testsDefinition and Common tests

Neuropsychological evidenceNeuropsychological evidence

Neuroimaging evidenceNeuroimaging evidence

SummarySummary


3/35

Taxonomy of MemoryTaxonomy of Memory

Memory

Declarative Non-declarative

Cohen and Squire, 1980


4/35


Memory


Available to conscious retrieval

Can be declared (propositional)

Examples What did I eat for breakfast?

(episodic) What is the capital of Spain?

(semantic) What did I just say?

(working)

Experience-induced change inbehaviour

Cannot be declared (procedural)

Examples Subliminal advertising?

(priming) How to ride a bicycle

(skills) Phobias

(conditioning)

Cohen and Squire, 1980


5/35


Memory


Semantic WorkingEpisodic


6/35


Memory




7/35

Episodic MemoryEpisodic Memory

Memory for personally experienced events that occurred in particularMemory for personally experienced events that occurred in particular

placeplace

at a specificat a specific

timetime (defined by Tulving, 1972)(defined by Tulving, 1972)

Contextual, spatiotemporal, autobiographical, rememberingContextual, spatiotemporal, autobiographical, remembering

Direct memory tests:Direct memory tests:

EncodingEncoding RetrievalRetrieval

Free recallFree recall CATCAT ??

DOGDOG ??

Cued recallCued recall CAT DOGCAT DOG EAGLE- ?EAGLE- ?

EAGLE NESTEAGLE NEST CAT- ?CAT- ?

RecognitionRecognition CATCAT CATCAT

DOGDOG SUNSUN XX

Source MemorySource Memory CATAT CATCAT boldboldDOGDOG DOGDOG italicsitalics


8/35

Episodic MemoryEpisodic Memory

Copy

DelayedRecall(15 mins)

Rempel-Clower et al., 1996

CONTROLSPATIENTS


9/35

Episodic Memory - NeuropsychologyEpisodic Memory - Neuropsychology

Organic AmnesiaOrganic Amnesia Intact:Intact: semantic memory (e.g, language)semantic memory (e.g, language)

working memory (e.g,


10/35

Large lesions of bilateral Hippocampi, Amygdalae, and Rhinal Cortex producesevere antero- and retro-grade amnesia, eg, HM (Scoville & Milner, 1957)

Circumscribed lesions of CA1 of Hippocampus produce significant anterogradeamnesia (Zola-Morgan et al 1986)

Korsakoffs Patients with diffuse damage to Diencephalon, Medial Thalamus,Mammillary Nuclei show varied amnesia (Press et al., 1989)

Alzheimers patients show early signs of amnesia, with first lesions in MedialTemporal Lobe (Hyman et al 1984)

Frontal Patients show confabulation (Burgess & Shallice, 1996), impaired sourcememory (Janowsky et al., 1989) and interference (Shimamura et al., 1995)

Episodic Memory - NeuropsychologyEpisodic Memory - Neuropsychology


11/35

MTL activations during episodic encoding (Tulving et al 1996) and retrieval (Schacter

et al. 1996)

Anterior-Posterior dissociation? (Lepage et al. 1998; Schacter et al. 1999)

Left Frontal during Encoding (Shallice et al., 1994), right during Retrieval

HERA: Hemispheric Encoding Retrieval Asymmetry (Tulving et al., 1994)

Posterior cingulate / Precuneus (Fletcher et al., 1996)

Left lateral inferior parietal cortex (Henson et al., 1999)

Network of Frontal - Medial Temporal Posterior areas all involved:

Frontal areas control encoding and retrieval of memories?Posterior association areas store components of memories?

Medial Temporal regions (temporarily) bind different components?

Finer spatial resolution (fMRI) beginning to dissociate MTL regions, egHippocampus / Perirhinal for Recollection / familiarity? (Aggleton & Brown, 1999)

Episodic Memory - NeuroimagingEpisodic Memory - Neuroimaging


12/35


Memory




13/35

Semantic MemorySemantic Memory Memory for facts, general knowledge, word meaningsMemory for facts, general knowledge, word meanings

Acontextual: Independent of where or when the information was encodedAcontextual: Independent of where or when the information was encoded

Pyramid and Palm Tree Test(Howard & Patterson 1992)

Common Tests:Common Tests:

Object NamingObject Naming e.g. What is this?e.g. What is this?

Semantic JudgementsSemantic Judgements

Which bottom picture goes best with the top one?Which bottom picture goes best with the top one?

Category FluencyCategory Fluency

Name as many dog breeds as possible in 1 minuteName as many dog breeds as possible in 1 minute

German Shepard, golden retriever, . . .German Shepard, golden retriever, . . .


14/35

Modality-specific visual Agnosia after Left Temporal damage (Warrington, 1975)

Category-specific, amodal Agnosia following Left Medial/Middle Temporal damage,eg, living vs. nonliving (Warrington & Shallice, 1984)

category effects reflect visual vs functional information?

Temporal Pole lesions cause deficits in person-naming; Left Middle Inferior inanimal-naming and Left Posterior Inferior in tool-naming (Damasio et al 1996)

Semantic Dementia (SD) following anterior/inferior Temporal atrophy, with reverseRibot gradients (Graham et al., 2000)

SD patients demonstrate graded deterioration of knowledge (Hodges et al., 1992)

Semantic Memory - NeuropsychologySemantic Memory - Neuropsychology


15/35

Common activation in Left Inferior Frontal,Inferior Temporal, Angular gyrus and Temporalpole for semantic judgments to words andpictures (Vandenberghe et al 1996)

Left Inferior Temporal activations for animal

and tool naming, Temporal Pole for peoplenaming (Damasio et al., 1996)

Left Inferior Temporal activation for category-versus letter-fluency (Mummery et al 1996)

Left Middle Temporal and Premotor activationsfor tool vs animal naming, Left Middle Occipitalfor animal vs tool naming (Martin et al 1996)

Distributed representations, with activationsreflecting objects interaction with world? E.g.,

tool naming activates motor regions

Semantic Memory - NeuroimagingSemantic Memory - Neuroimaging

McClelland and Rogers, 2003


16/35

Are episodic/semantic memory just a continuum? (Watkins, 1974)

Does Global Anterograde Amnesia exist?

Inability to acquire new semantic memories (Gabrieli et al, 1998)...

...yet intact development of semantic memory despitehippocampal damage (Vargha-Khadem et al. 1998)

Hippocampus proper underlies true episodic memory?

Additional Medial Temporal areas underlie anterograde semanticmemory impairment?

Episodic vs Semantic debateEpisodic vs Semantic debate


17/35


Memory




18/35

Common TestsCommon Tests

Memory Span (maintenance)Memory Span (maintenance)

Digit span Repeat back: 8,5,3,2,7,9Digit span Repeat back: 8,5,3,2,7,9

Working MemoryWorking Memory The ability to hold information on-line for current task (e.g. for comprehension,The ability to hold information on-line for current task (e.g. for comprehension,

planning, problem solving, Baddeley 1992)planning, problem solving, Baddeley 1992)

Short-term memory (cf. long-term episodic / semantic memory)Short-term memory (cf. long-term episodic / semantic memory)

Verbal vs VisuospatialVerbal vs Visuospatial Maintenance vs ManipulationMaintenance vs Manipulation Spatial vs ObjectSpatial vs Object Storage vs RehearsalStorage vs Rehearsal

Corsi Block spanCorsi Block span

N-back task (manipulation)N-back task (manipulation)

. . . $ % ^ ! * & *. . . $ % ^ ! * & * +

(Sternberg) probe task (maintenance)(Sternberg) probe task (maintenance)


19/35

Auditory-Verbal maintenance deficit following Left Supramarginal / InferiorParietal lesions, eg KF (Warrington & Shallice, 1969)

Visual-spatial maintenance deficit following Right Inferior Parietal lesion, eg ELD(Hanley et al 1991)

Frontal patients impaired on manipulating information in Working Memory ontasks such as card sorting (Milner, 1963) and selection-without-repetition(Petrides & Milner, 1982)

Age-related Working Memory deficits following frontal-striatal decline (Gabrieli,1996)

Modality-specific, passive stores in posterior parietal/temporal cortex

Common executive processes in frontal cortex

Working Memory - NeuropsychologyWorking Memory - Neuropsychology


20/35

Left Inferior Parietal activation during verbal storage; Left Inferior Frontal andPremotor activation during verbal rehearsal (Paulesu et al. 1993)

Right Inferior Parietal, Inferior Frontal, Anterior Occipital, and PremotorCortex activated during spatial maintenance (Jonides et al. 1993)

Left inferior temporal and inferior parietal activated when object compared tospatial maintenance (Smith et al. 1995)

Dorsolateral Frontal Cortex activated in N-back task when manipulationrequired by N>1 (Cohen et al 1997), for both verbal and spatial material(Owen et al 1998)

Ventrolateral (Inferior) Frontal Cortex involved in maintaininginformation on-line in current form

Dorsolateral Frontal Cortex involved in manipulating information into

new forms (Owen 1997; Postle & DEsposito, 1999)

Working Memory - NeuroimagingWorking Memory - Neuroimaging


21/35


Memory


Semantic WorkingEpisodic Priming Procedural Conditioning


22/35


Memory




23/35

Priming A change in speed, accuracy or bias of processing aA change in speed, accuracy or bias of processing a

stimulus owing to prior exposure to that stimulusstimulus owing to prior exposure to that stimulus

Perceptual vs ConceptualPerceptual vs Conceptual

Example Indirect Memory Tests:Example Indirect Memory Tests:

Perceptual Identification Gollin Figures ->Perceptual Identification Gollin Figures ->

Perceptual Priming

(Gollin Figures)

Warrington & Weiskrantz, 1970

Stem/Fragment CompletionStem/Fragment Completion

(SMILE)(SMILE) SMI_ _SMI_ _

S_ _ L _S_ _ L _

Word AssociationWord Association

(ROSE)(ROSE) FLOWER - ?FLOWER - ?

Speeded DecisionsSpeeded Decisions

(APPLE)(APPLE) Concrete/Abstract?Concrete/Abstract?


24/35

Priming - Neuropsychology

Amnesics with Medial Temporal damage show intact PerceptualAmnesics with Medial Temporal damage show intact PerceptualPriming (Warrington & Weiskrantz 1970) and intact ConceptualPriming (Warrington & Weiskrantz 1970) and intact ConceptualPriming (Graf et al 1985)Priming (Graf et al 1985)

Huntingtons patients with Basal Ganglia damage show intactHuntingtons patients with Basal Ganglia damage show intact

priming (Heindel et al 1989)priming (Heindel et al 1989)

Alzheimers patients with diffuse Temporal Lobe damage showAlzheimers patients with diffuse Temporal Lobe damage showintact perceptual priming but impaired conceptual priming (Keaneintact perceptual priming but impaired conceptual priming (Keaneet al 1995)et al 1995)

Patients with right occipital lesions show no perceptual priming,Patients with right occipital lesions show no perceptual priming,but intact conceptual priming (Gabrieli et al 1995)but intact conceptual priming (Gabrieli et al 1995)


25/35

Priming - Neuroimaging

Reduced activity in bilateral occipito-temporal regions in word-stem completionReduced activity in bilateral occipito-temporal regions in word-stem completion

(Buckner et al. 1995), independent of explicit memory (Schott et al, 2005)(Buckner et al. 1995), independent of explicit memory (Schott et al, 2005)

Reduced activity in left frontal cortex in word-association (Blaxton et al 1996)Reduced activity in left frontal cortex in word-association (Blaxton et al 1996)

Subliminal priming right thru to motor cortex (Dehaene et al, 2001); thoughSubliminal priming right thru to motor cortex (Dehaene et al, 2001); though

issues of stimulus vs response priming (Dobbins et al, 2004)issues of stimulus vs response priming (Dobbins et al, 2004)

Left frontal cortex involved in conceptual/semantic primingLeft frontal cortex involved in conceptual/semantic priming

Occipito-temporal cortex involved in visual perceptual primingOccipito-temporal cortex involved in visual perceptual priming

Priming deactivations localised in same areas that performed initialPriming deactivations localised in same areas that performed initialprocessing (Schacter & Buckner, 1998)processing (Schacter & Buckner, 1998)

Deactivations reflect less neural activity (lowered thresholds, synapticDeactivations reflect less neural activity (lowered thresholds, synapticchange, residual activation)?change, residual activation)?Priming-related increases? (Henson, 2003)Priming-related increases? (Henson, 2003)


26/35


Memory




27/35

Procedural MemoryProcedural Memory

Skill learning (e.g. riding a bicycle)Skill learning (e.g. riding a bicycle)

Requires multiple trialsRequires multiple trials

Indexed by improved accuracy or RTsIndexed by improved accuracy or RTs

Mirror TracingMirror Tracing(e.g. Corkin, 1968)(e.g. Corkin, 1968)

RotaryPursuitRotaryPursuit(e.g. Gabrieli et al., 1997)(e.g. Gabrieli et al., 1997)

Serial Reaction TaskSerial Reaction Task

(e.g. Hazeltine et al., 1997)(e.g. Hazeltine et al., 1997)


28/35

Procedural - Neuropsychology

Amnesic patients show intact:Amnesic patients show intact: Rotary Pursuit (Corkin 1968)Rotary Pursuit (Corkin 1968) Serial Reaction Task (Nissen & Bullemer 1987)Serial Reaction Task (Nissen & Bullemer 1987)

Alzheimers patients show intact:Alzheimers patients show intact: Rotary Pursuit (Gabrieli et al 1993)Rotary Pursuit (Gabrieli et al 1993) Mirror Tracing (Heindel et al 1989)Mirror Tracing (Heindel et al 1989)

Parkinsons patients impaired on:Parkinsons patients impaired on: Rotary Pursuit (Heindel et al 1989)Rotary Pursuit (Heindel et al 1989) Serial Reaction Task (Ferraro et al 1993)Serial Reaction Task (Ferraro et al 1993)

Huntingtons patients impaired on:Huntingtons patients impaired on: Rotary Pursuit (Gabrieli et al 1997)Rotary Pursuit (Gabrieli et al 1997) Serial Reaction Task (Willingham & Koroshetz 1993)Serial Reaction Task (Willingham & Koroshetz 1993)

but not:but not: Mirror Tracing (Gabrieli et al 1997)Mirror Tracing (Gabrieli et al 1997)

Cerebellar lesions impair Mirror Tracing (Sanes et al 1990)Cerebellar lesions impair Mirror Tracing (Sanes et al 1990)


29/35

Procedural - Neuroimaging Rotary Pursuit learning correlates with activity in Primary and Secondary MotorRotary Pursuit learning correlates with activity in Primary and Secondary Motor

Cortex (Grafton et al 1992)Cortex (Grafton et al 1992)

Serial Reaction Task correlates with activity in Primary and Secondary MotorSerial Reaction Task correlates with activity in Primary and Secondary MotorCortex, and Basal Ganglia (Hazeltine et al 1997)Cortex, and Basal Ganglia (Hazeltine et al 1997)

Two hypotheses:Two hypotheses:

1.1. Learning repetitive sequence involves Basal Ganglia-Thalamic-Motor Cortical loopLearning repetitive sequence involves Basal Ganglia-Thalamic-Motor Cortical loop

Learning new visual-motor mappings involves Cerebellar-Motor Cortical loopLearning new visual-motor mappings involves Cerebellar-Motor Cortical loop

2.2. Open-loop learning (minimal feedback): Basal Ganglia-Thalamic-Motor Cortical loopOpen-loop learning (minimal feedback): Basal Ganglia-Thalamic-Motor Cortical loop

Closed-loop learning (continual feedback): Cerebellar-Motor Cortical loopClosed-loop learning (continual feedback): Cerebellar-Motor Cortical loop

Rotary Pursuit and Serial Reaction Task involve open-loop motor learning with littleRotary Pursuit and Serial Reaction Task involve open-loop motor learning with littlevisual feedback (impaired by Basal Ganglia lesions)visual feedback (impaired by Basal Ganglia lesions)

Mirror Tracing involves much visual feedback (impaired by Cerebellar lesions)Mirror Tracing involves much visual feedback (impaired by Cerebellar lesions)

Need to examine nonvisual feedbackNeed to examine nonvisual feedback


30/35


Memory




31/35

Classical ConditioningClassical Conditioning Changes in response (R) to conditioned stimulus (CS) after repeatedChanges in response (R) to conditioned stimulus (CS) after repeated

conditioned unconditioned stimulus (US) pairingconditioned unconditioned stimulus (US) pairing

Example:Example:

Existing:Existing: e.g. air puff to eye (US) blink reflex (R)e.g. air puff to eye (US) blink reflex (R)

Training:Training: e.g. tone in ear (CS) air puff to eye (US)e.g. tone in ear (CS) air puff to eye (US)

Result:Result: tone in ear (CS) blink reflex (R)tone in ear (CS) blink reflex (R)

Delay Conditioning: US starts after a CS but they co-occurDelay Conditioning: US starts after a CS but they co-occur

CS

US

CS

US

Trace Conditioning: US starts after CS but they do NOT co-occurTrace Conditioning: US starts after CS but they do NOT co-occur

Fear Conditioning: CS is neutral (e.g. a light), US is aversive (e.g. shock)Fear Conditioning: CS is neutral (e.g. a light), US is aversive (e.g. shock)

R is behavioural/physiological (e.g. Galvanic skin response)R is behavioural/physiological (e.g. Galvanic skin response)


32/35

Conditioning - Neuropsychology Delay conditioning (eye-blink):Delay conditioning (eye-blink):

Abolished with Cerebellar lesions (Daum et al 1993)Abolished with Cerebellar lesions (Daum et al 1993)

Abolished in Alzheimers Disease: diffuse damage? (Woodruff-Pak et al 1990)Abolished in Alzheimers Disease: diffuse damage? (Woodruff-Pak et al 1990)

Impaired in Korsakoffs amnesics: diffuse damage (McGlinchey-Berroth et al 1995)Impaired in Korsakoffs amnesics: diffuse damage (McGlinchey-Berroth et al 1995)

Intact despite Basal-Ganglia lesions in Huntingtons (Woodruff-Pak & Papka 1996)Intact despite Basal-Ganglia lesions in Huntingtons (Woodruff-Pak & Papka 1996)

Intact in Medial-Temporal amnesics (Gabrieli et al 1995b; Clark & Squire, 1998)Intact in Medial-Temporal amnesics (Gabrieli et al 1995b; Clark & Squire, 1998)

Trace conditioning:Trace conditioning:

Impaired in Medial-Temporal amnesics (McGlinchey-Berroth et al 1997)Impaired in Medial-Temporal amnesics (McGlinchey-Berroth et al 1997)

Fear conditioning:Fear conditioning:

Impaired following amygdala resection, despite intact declarative memory forImpaired following amygdala resection, despite intact declarative memory for

contingency (LeBar et al 1995)contingency (LeBar et al 1995)

Intact in amnesics despite impaired declarative memory for contingencyIntact in amnesics despite impaired declarative memory for contingency

(Bechara et al 1995)(Bechara et al 1995)


33/35

Conditioning - Neuroimaging Cerebellar activity correlated with CR during Delay ConditioningCerebellar activity correlated with CR during Delay Conditioning

(Logan & Grafton 1995)(Logan & Grafton 1995)

Hippocampus (and amygdala) shows transient involvement in FearHippocampus (and amygdala) shows transient involvement in Fear

Trace Conditioning (Buechel et al., 1999)Trace Conditioning (Buechel et al., 1999)

Amygdala activity correlates with CS during Fear conditioning (MorrisAmygdala activity correlates with CS during Fear conditioning (Morris

et al 1998)et al 1998)

Cerebellum implicated in delay conditioningCerebellum implicated in delay conditioning

Amygdala implicated in fear conditioningAmygdala implicated in fear conditioning

Hippocampus may be involved in trace conditioningHippocampus may be involved in trace conditioning

(development of declarative memory for contingency?)(development of declarative memory for contingency?)

Future research may benefit from analyses of effectiveFuture research may benefit from analyses of effective

connectivity (Buechel et al 1998)connectivity (Buechel et al 1998)


34/35


Memory



What did Ijust say?

What did Ihave for

breakfast?

What is thecapital of

France?

Facilitatedprocessing

Reflex responseto new stimuli

Lateral FrontalParietal and

occipitalMedial temporalDiencephalon

Mammillary bodiesFrontal lobe

LateralTemporal /

Frontal lobes

Many corticalregions

Cerebellum/Amygdala

(MTL?)

How to ridea bicycle

Basal GangliaCerebellum

Motor cortex


35/35

ReferencesBaddeley (1992) Science, 255, 556-559Bechara et al (1995). Science, 269, 1115-1118Blaxton et al (1996) Can. J. Exp. Psych., 50, 42-56Buckner et al (1995) J. Neurosci., 15, 12-29Buechel et al. (1999) J. Neurosci., 19, 10869-10876Burgess & Shallice (1986) Memory, 4, 359-441

Clark & Squire (1998). Science, 280, 77-81.Cohen & Squire (1980), Science, 210, 207-210Corkin (1968) Neuropsychologia, 6, 255-265Damasio et al (1996) Nature, 380, 499-505Daum et al (1993) J. Neurol. Neurosurg. Psychiatry, 52, 47-51Ferraro et al (1993) Brain Cog., 21, 163-180Fletcher et al (1996) Brain, 119, 1587-1596Gabrieli et al (1993) Behav. Neurosci., 107, 899-910Gabrieli et al (1995) Psych. Sci, 6, 76-82Gabrieli et al (1995b) Behav. Neurosci., 109, 819-827Gabrieli (1996) Proc. Nat. Acad. Sci. USA, 93, 13534-13540Gabrieli et al (1997) Neuropsychology, 11, 272-281Gabrieli et al (1998) Brain Cog., 7, 525-539Gabrieli, J. (1998). Annu. Rev. Psychol. 49: 87-115Graf et al (1985) J. Exp. Psych: Learn. Mem & Cog, 11, 501-518Grafton et al (1992) J. Neurosci., 12, 2542-2548Hanley et al (1991) Quart. J. Exp. Psych., 43A, 101-125

Hazeltine et al (1997) Brain, 120, 123-140Heindel et al (1989) J. Neurosci., 9, 582-587Henson et al (1999) J. Neurosci., 19, 3962-3972Hodges et al (1995) Brain, 15, 1783-1806Hyman et al (1984) Science, 225, 1168-1170Janowsky et al (1989) Behav. Neurosci., 103, 548-560Jonides et al (1993) Nature, 363, 623-625Keane et al (1997) Neuropsych., 11, 59-69LeBar et al (1995) J. Neurosci., 15, 6846-6855

LePage et al (1998) Hippocampus 8 313-322

Logan & Grafton (1995) Proc. Nat. Acad. Sci. USA, 92, 7500-7504McGlinchley-Berroth et al (1995) Alcohol Clin. Exp. Res., 1127-1132McGlinchley-Berroth et al (1997) Behav. Neurosci.Milner (1963) Arch. Neurol., 9, 90-100Morris et al (1998) Nature, 393, 467-470Martin et al (1996) Nature 379, 649-652

Mummery et al (1996) Proc. Roy. Soc. London B, 263, 989-995Owen (1997) Euro. J. Neurosci., 9, 1329-1339Owen et al (1998) Proc. Nat. Acad. Sci. USA, 95Nissen & Bullemer (1987) Cog. Psych., 19, 1-32Paulesu et al (1993) Nature, 362, 342-344Peterson et al (1989) J. Cog. Neurosci., 1, 153-170Petrides & Milner (1982) Neuropsychologia, 20, 601-614Press et al (1989) Nature, 341, 54-57Rugg (1995) Electrophysiology of mind. Oxford: OUP

Schacter et al (1996) Proc. Nat. Acad. Sci. USA, 93, 321-325Schacter & Buckner (1998) Neuron, 20, 185-195Scoville & Milner (1957) J. Neurol. Neurosurg. Psychiatry, 20, 11-21Shallice et al (1994) Nature, 368, 633-635Shimamura et al (1995) J. Cog. Neurosci., 7, 144-152Smith et al. (1995) J. Cog. Neurosci., 7, 337-356.Tulving (1983) Elements of episodic memory. London: OUPTulving et al (1994) Proc. Nat. Acad. Sci. USA, 91, 2016-2020

Vargha-Khadem et al (1997) Science, 277, 376-380Warrington (1975) Quart. J. Exp. Psych., 27, 635-657Warrington & Shallice (1984) Brain, 106, 859-878Warrington & Shallice (1969) Brain, 92, 885-896Warrington & Weiskrantz (1970) Nature, 228, 628-630Willingham & Koroshetz (1993) Psychobiology, 21, 173-182Woodruff-Pak et al (1990) Clin. Neurosci. Neuropath., 1, 45-49Woodruff-Pak & Papka (1996) J. Int. Neuropsychol. Soc, 2, 323-334Sanes et al (1990) Brain, 113, 103-120

Tulving et al (1996) Cerebral Cortex 6 71-79Z l M t l (1986) J N i 6 2950 2967

Cbu Human Memory

Documents

Transcript of Cbu Human Memory