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Prof. Stephan Anagnostaras
Lecture 6: Multiple Memory Systems: Implicit Memory
Neurobiology ofLearning and Memory
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Different types of learning & memory rely on different brainstructures
Explicit memoryImplicit memory
Facts(semantic)
Events(episodic)
Medial temporal lobe; diencephalon
Procedural memory:skills & habits(basal ganglia) Skeletal musculature
(cerebellum)
Classical conditioning
Emotional Responses(amygdala)
Priming(neocortex)
Eyeblink conditioning inrabbit
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Squire’s Taxonomy of Memory
Squire & Zola, PNAS, 1996
Implicit memory is a broaderterm than explicit memory
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Basal Ganglia• Can examine Parkinson’s & early Huntington’sDisease
• no apparent amnesia (declarative memory ok)
But implicit memory problems in“procedural memory”• Perceptual-Motor Learning• Habits• Skills
Separate from motor disorders
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Serial Reaction Time (SRT) Task
Subjects are not told about the sequence.
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SRT Results
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Several perceptual motor tasksimpaired by basal ganglia damage
• Serial Reaction Time
• Backwards Reading
• Prism adaptation
• Mirror drawing
• Also more cognitive tasks…
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Knowlton, Mangels, & Squire, Science, 1996
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Artificial Grammar Learning
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Prototype Abstraction
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Basal ganglia
• Implicit learning deficits in several tasks
• Don’t need a motor component
• Can be quite “cognitive”
• No explicit memory necessary
• What about animals?
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Packard, Hirsch & White,J Neurosci 1989
McDonald & White, BehNeurosci 1993
% correct
Time spent
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Win-Stay Acquisition Trials
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Double dissociation paper:Electrolytic lesions on win-stay
Packard, Hirsh & White, J Neurosci, 1989
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Win-Shift Training
Training Phase
Baited
Delay
Testing Phase
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Double dissociation paper:Electrolytic Lesions on win-shift
Packard, Hirsh & White, J Neurosci, 1989
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Triple D Histology (HPC + DLC)
McDonald & White, Behav Neurosci, 1993
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Triple D: Neurotoxic DLC lesionsimpair win-stay learning
McDonald & White, Behav Neurosci, 1993
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Triple D: Neurotoxic DLC lesions sparewin-shift learning
McDonald & White, Behav Neurosci, 1993
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Triple D: Neurotoxic HPC lesionsimpair win-shift learning
McDonald & White, Behav Neurosci, 1993
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Triple D: Neurotoxic BLA lesionsimpair conditioned cue preference
McDonald & White, Behav Neurosci, 1993
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Proposed characteristics of habitlearning
Squire, 1992:
• Knowledge expressed through performance, rather than recollection• Associations acquired across many trials• Less flexible (less transferable) than declarative learning
Salmon & Butters, 1995: “Habit learning refers to the formation of simple associations in which a neutral stimulus comes to elicit a certain motor response as a function of repeated reinforcement.”
Mishkin et al., 1984:Product of processing is “not cognitive information, but a non-cognitive simulus-response (S-R) bond…what is stored is not objects, emotions…but the changing probability that a given stimuluswill evoke a specific response due to the reinforcement contingencies at that time”
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Sage & Knowlton, Beh Neurosci, 2000US Devaluation of Win-Stay
US Devaluation(Testing the hypotheses)
Protocol: Train
US devaluation
Test
• If performance does not involve representation of US, the CR should remain intact (e.g., accurate, fast) on test. This would be consistent with an S-R view.
• If performance mediated by representation of US, then recall of devalued US should change the CR (e.g., inaccurate, slow) on test
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Devaluation following win-stay acquisition
ControlUnpairedPaired
Trial
Corr
ect (
%)
0
20
40
60
80
100
1 5 10 15 20 22 Probe5
Acquisition
5
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0
20
40
60
80
100
Corr
ect (
%)
1 2 3 4 5 6 7 8 Trials within Post-CTA
Extinction Probe
A. Extinction Accuracy
ControlUnpairedPaired
Win-stay late timepoint post-CTA trials
xx
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Post-CTA win-stay accuracy summary
Timepoint(Amount of Training)
2, 9, or 22 days
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Post-CTA win-stay latency summary
Timepoint(Amount of Training)
* = significant difference
* *
*
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Contextual Cuing Task(Chun & Phelps, Nat Neurosci, 1999)
Easy Version Difficult Version
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Contextual Cuing Task(Chun & Phelps, Nat Neurosci,1999)
Implicit Learning Deficit in Amnesia !
Amnesics2 anoxic (1 conf hipp)2 encephalitic (2 conf hipp)
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Memory systems in Implicit Memory
• Not all implicit memory is independent ofthe hippocampus
• Not all implicit memory depends on thebasal ganglia, e.g., emotional learning,priming, certain motor responses
• Cortical systems (e.g., priming)• Amygdala (fear conditioning)• Cerebellum (eyeblink conditioning)…etc