PS4529/30 Applications of Cognitive Neuroscience
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Transcript of PS4529/30 Applications of Cognitive Neuroscience
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PS4529/30PS4529/30ApplicationsApplicationsof Cognitiveof CognitiveNeuroscienceNeuroscience
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Consensus View of Long-Term Memory
ENCODING RETRIEVAL
Seeing Word
Hearing Word
MTL
MTL
MTL
MTL
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A Specific Example
• The constructive memory framework (CMF)• Schacter, DL, Norman, KA, and Koutstaal, W.
(1998). The cognitive neuroscience of constructive memory. Annual Review of Psychology, 49, 289-318.
• Invokes multiple brain regions• Some involved in encoding and retrieval• Some involved in either encoding or retrieval
• Comprising multiple functions that must interact dynamically with one another
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CMF Neuroanatomy
• The hippocampal formation ‘Indexing’ of episodes: exactly how is unknown Necessary both for encoding and retrieval Damage leads to dense retrograde and anterograde
amnesia
• The frontal lobes Strategic control over memory: exactly how is again
unknown! Damage leads to confabulations, delusions, heightened
false memory, source amnesia
• The entire ‘association’ neocortex Representation of experienced content Damage should lead to loss of specific content of prior
episodes
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CMF Retrieval Functions
Retrieval ‘focus’
Access to the records of attended information via a retrieval cue (by hippocampal pattern completion)
Inhibition of irrelevant information
Re-activation of episodic content (held in the neocortex)
Monitoring/evaluating retrieval products (prefrontally mediated)
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How Many Experiences Have you Had?
1
10
100
1000
10000
100000
1000000
10000000
100000000
1000000000
HOUR DAY WEEK MONTH YEAR DECADE
TIME
Num
ber
of e
piso
des
(log)
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Conway’s SMS Model
The Self Memory System (SMS) has two principle components:-
• 1. Autobiographical knowledge base– organised specifically to support our
sense of self
• 2. The (working) Self– comprises a goal hierarchy, and various
other internal mechanisms
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Autobiographical Knowledge Base
‘Self’
Episodic Memory tied to specific experiences
(e.g. the CMF)
Retrieval
Consolidation
Encoding
Self-relatedsemantic knowledge
Abstracted from specificexperiences
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Goal Hierarchy
Take a peek inside yourself…
Conceptual Self
Self
Autobiographical Knowledge Base
semantic knowledge
Episodic Memory(CMF)
Key point: the SMS system is ‘goal-driven’
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The SMS greatly extends the CMF
• Conway: “all daily experiences are destined to be forgotten”– Unless they support longer-term goals
• In the short term, accurate memories are vital– Where did I leave my keys
• In the long-term, coherence (between goals) is vital– The ‘Husband-Hermit’ or ‘Saint-Sinner’ dilemma
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SMS Goals• Short term (e.g. daily)
– Take the car for a service…– Find the keys…– Post the letter…
Overriding principle: accuracy!
• Long-term– Get a job– Learn how to drive– Buy a house– Become a solitary religious hermit– Be a loving husband
• Overriding principle: coherence
But: there is an accuracy-coherence trade-off!
Potential for conflict!
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How is the trade-off achieved?
• The goal hierarchy maintains a stable and coherent set of short and long term goals
Goal Hierarchy
Eat and drink (everyday)Keep warm (everyday)
Have a conversation (most days)Watch TV (particular times)
Find the car keys (in 5 minutes)Avoid tripping up (when I walk)
Post the letter (sometime today)Dentist appointment (this week)
Revise (next month)Obtain graduation ball tickets (next few
months)Find a less annoying partner (yesterday!)
Get a 2:1 (next couple of years)Loose weight (before going on holiday)
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The SMS: key points
• New memories are not formed ‘automatically’ from our experiences– But, experiences are always encoded (e.g. as per the CMF).
• An ABM is formed (and retrieved) only when the (working) self interacts with the autobiographical knowledge base
– Such interactions are entirely goal-driven
• Hence, specific experiences will be forgotten unless they relevant to a goal - within the goal hierarchy
• Stable self-image emerges from the coherence imposed by the goal hierarchy, perhaps at the expense of accuracy
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Autobiographical Knowledge Base
‘Self’
Episodic Memory tied to specific experiences
(e.g. the CMF)
Retrieval
Consolidation
Encoding
Self-relatedsemantic knowledge
Abstracted from specificexperiences
Is this: (1) A scientifically acceptable and (2) A forensically useful model?
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Can ERPs reveal exactly what is happening in the brain while people remember their past?
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cueonset
Ecphory/inhibition
MonitoringRetrieval Perception/attention
Patterncompletion/
Binding
‘selective attention’
Stimuli
Time 0.1 0.2 0.40.3 0.5 0.70 0.6
Constructive Memory Framework
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Focussed Search
Retrieve / Inhibit Monitor
Stimuli
0Time - 1 3 4 65 7 92 8
Retrieval success!!Retrieval failure
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ERP correlates of retrieval from long-term memory
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Stimuli
Time 0.1 0.2 0.40.3 0.5 0.70 0.6
Ecphory?
Monitoring?
Implicit Memory?
Familiarity?
Donaldson, Allan and Wilding (2003)Mecklinger (2000)Rugg and Wilding (2000)
F-N400
LP effect
RF effect
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Content only (versus failure)
Content and Context
LP effect magnitude X content relation
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Using ERPs to investigate the notion of encoding-retrieval overlap
1. By manipulating the content of what is encoded and retrieved.
2. By manipulating the timing of encoding and retrieval, to make them coincide.
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1. Manipulating Content
• Operationally define different classes of study episode
• Record EEG when instances of each class of episode are recollected
• Form ERPs to each class of recollected episode
• Contrast the magnitude and topography of ERPs for each class of recollected episode
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Encoding and Retrieval in vivo…
Olfactory (Gottfried et al, 2004)
and within ‘sensory domain’ too (Woodruff et al., 2005)
MTL
Encoding
MTL
Retrieval
TIMEVisual
Auditory
MTL
Encoding
MTL
Retrieval
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Do ERPs revealDo ERPs revealmodality specific retrieval processes?modality specific retrieval processes?
• Subjects SAW and HEARD words at study
• Performed a word-stem (e.g. MOT__) cued recall task
• ERPs were formed to stems completed with Studied SEEN items Studied HEARD items Unstudied NEW items
• ERP retrieval effects for each sensory modality:- SEEN – NEW difference HEARD – NEW difference
Allan, Robb and Rugg (2000), Neuropsychologia, 38 1188-1205.
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No!ERPs are insensitive to
differences in modality at retrieval
Recall auditory episodeRecall visual episode
As retrieval ends…
As retrievalbegins…
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ERP Modality Experiment: ConclusionsERP Modality Experiment: Conclusions
• Multiple retrieval processes, active at different times– Onset ~ 0.5s after retrieval cue!
• Retrieval of ‘visual’ and ‘auditory’ episodes involves common processes. No evidence for modality specific retrieval processes.
• ERPs reflect a ‘core component’ of retrieval? – Changes in neocortical activity driven by the
Hippocampus during early stages of retrieval (prior to modality specific activations)?
– Or: attention to retrieval products?
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Episodic Memory Mechanisms
ConsolidationMechanisms
AttentionalControl
Encoding Storage Retrieval
AttentionalControl
SemanticRecords
PerceptualRecords
Binding
ContextSemanticRecords
PerceptualRecords
Binding
Context
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• Gain precise control over the relative timing of events experienced in different modalities.
• Stress the system by forcing it to handle very rapidly changing inputs, to reveal what the temporal limits are.
• Examine resulting performance behaviourally
• And use high temporal resolution neurophysiological data to expose the underlying functional states
2. Can we simultaneously encode and retrieve?
Allan and Allen (2005), Journal of Neuroscience, 25, 8122-9130.
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Does encoding temporarily stop when retrieval occurs?
How Many Experiences Have you Had?
1
10
100
1000
10000
100000
1000000
10000000
100000000
1000000000
HOUR DAY WEEK MONTH YEAR DECADE
TIME
Num
ber
of e
piso
des
(log)
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Visual
Auditory
+/- 200msec period of stimulus onset asynchrony (SOA) jitter, in 10 time bins (41 – 194msec)
E == encode (animacy task)
R == retrieve (old/new recognition)
Key
Phase 1
Phase 2
Phase 3
E E E
R
E
R
E
R R R
3-Phase Dual-task Paradigm3-Phase Dual-task Paradigm
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Stimulus-Onset Asynchrony (SOA)
Encode time-line begins
WILD
Retrieve time-line begins
Time controlled ‘jitter’
Expt. 1 SOA range: 50-200msecExpt. 2 SOA range: 50-2000msec
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0
10
20
30
40
50
60
70
80
% c
orr
ec
t
Full Hit 71.85 62.66
DA Hit 72.03 49.38
Full CR 79.21 71.04
DA CR 76.88
phase 2 phase 3
Dual-task Performance (Expt. 1)Dual-task Performance (Expt. 1)
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Effect of ‘ignoring’ retrieval cues
Retrieval under full attention
Retrieval under distractedattention
LP effect minimal/absent under DA, replaced by F-N400but no reliable topographic differences (Allan and Allen, 2005)
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• Encoding stabilised at a temporal gap of ~600msec(see Expt. 2), i.e. just as the ERP effect begins.
– retrieval cue processing is complete.– neocortical trace reactivation has commenced.– so ‘automatic’ encoding of experience can begin again?
• Retrieval shows a subtle alteration towards reliance on familiarity
• Mode-shifting between encoding and retrieval in human memory is relatively sluggish
•The LP effect may reflect the attention paid to retrieval products, not the representational nature of those products
Conclusions
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Next week -
Do these ERP effects objectively indicate the presence / absence of an episodic memory?