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Chapter 2:
Studies of Human
Learning and Memory
From Mechanisms of Memory, second edition
By J. David Sweatt, Ph.D.
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Learning: The acquisition of an altered behavioral
response due to an environmental
stimulus.
Memory: The processes through which learned
information is stored.
Recall: The conscious or unconscious retrieval
process through which this
altered behavior is manifest.
Definitions
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Movies Using Amnesia as a Plot Device
Movie Director (plus Notable Actors) YearAnterograde Amnesia:
Memento Christopher Nolan (Guy Pearce) 2001
Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003
50 First Dates Peter Segal (Adam Sandler) 2004
Retrograde Amnesia:
The Bourne Identity Doug Liman (Matt Damon) 2002
Total Recall Paul Verhoeven (Arnold Schwarzenegger, Sharon Stone) 1990
Spellbound Alfred Hitchcock (Ingrid Bergman, Gregory Peck) 1945
Eternal Sunshine Michel Gondry (Jim Carrey, Kate Winslet) 2004
of the Spotless Mind
The Forgotten Joseph Ruben (Julianne Moore) 2004
Men in Black Barry Sonnenfeld (Tommy Lee Jones, Will Smith) 1997
Not Sure:
Mulholland Drive David Lynch (Michael J. Anderson, Diane Baker) 2001
Table 1
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HUMAN MEMORY
DECLARATIVE
(EXPLICIT)
NONDECLARATIVE
(IMPLICIT)
FACTS EVENTS
PROCEDURAL
(SKILLS AND
HABITS)
PRIMING SIMPLE
CLASSICAL
CONDITIONING
NONASSOCIATIVE
LEARNING
EMOTIONAL
RESPONSES
SKELETAL
MUSCULATURE
MEDIAL
TEMPORAL
LOBE
STRIATUM NEOCORTEX AMYGDALA CEREBELLUM REFLEX
PATHWAYS
Subdivisions of Human Memory
Milner B, Squire LR, Kandel ER: "Cognitive neuroscience and the study of memory". Neuron 1998, 20:445-468.
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Multiple Memory Systems in the Human CNS
Memory Subtype Corresponding CNS Subregion
Working Memory Prefrontal Cortex, contributions from Caudate Nucleus
Declarative, Episodic, and Spatial Medial Temporal Lobe system including the
Memory Hippocampus, Dentate Gyrus, Ento-rhinal and Peri-
rhinal cortices
Habits, Motor Skills, Procedural Striatum (Caudate Nucleus and Putamen), Globus
Memory Pallidus*, and Cerebellum
Priming Occipital Cortex, Neocortex in general
Aversive Associative Conditioning Amygdala
Motoric Associative Conditioning Cerebellum
Olfactory and Taste Conditioning Olfactory Bulb, Insular Cortex, Nucleus Tractus
Solitarius (NTS), positive and negative reinforcement
systems (Amygdala or Nucleus Accumbens)
Sensitization, Habituation Spinal Cord, Brainstem Nuclei, Amygdala
Circadian Rhythm Hypothalamus – Suprachiasmatic Nucleus (SCN)
Reward, Positive Reinforcement, Nucleus Accumbens, Ventral Tegmental Area (VTA)
Addiction
*Collectively the Caudate Nucleus, Putamen, and Globus Pallidus are referred to as the Basal
Ganglia.
Table 2
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Memory-associated Cognitive Dysfunction in Humans
Disease or Syndrome Type of Memory Affected Likely Anatomical Locus
or Involved*
Schizophrenia Working Memory Prefrontal Cortex
Major Depression Declarative Memory ?
Aging-related Dementias Long-term Declarative and Temporal Lobe Hippocampal
Episodic Memory System
Korsakoff’s Syndrome Long-term Declarative and Mamillary Bodies, Limbic
(Alcoholism, Malnutrition) Episodic Memory System, Thalamus
Huntington’s Disease and Motor Learning, Habit Learning Nigro-striatal System
Parkinson’s Disease
Attention Deficit Generalized or Specific Learning Frontal Lobes?
Hyperactivity Disorder (ADHD) Disabilities, Dyslexia Basal Ganglia?
Post-traumatic Aversive Associative Conditioning, Amygdala, Hippocampus
Stress Disorder* (PTSD) Lack of Extinction
Table 3
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Figure 6A
Short-term Memory and the Multi-store Memory Model
CONTROL PROCESSES
•Stimulus analyzer programs
•Alter biases of sensory channels
•Activate rehearsal mechanism
•Modify information flow from SR to STS
•Code and transfer information from STS to LTS
•Initiate or modify search of LTS
•Heuristic operations on stored information
•Set decision criteria
•Initiate response generator
RESPONSE
GENERATOR
S
E
N R
S E
O G
R I
Y S
T
E
R
STIMULUS
INPUT
RESPONSE
OUTPUT
REHEARSAL
BUFFER
SHORT-TERM
STORE
Memory bank subject
to rapid decay
LONG-TERM
STORE
Self-addressable memory bank
not subject to decay
REHEARSAL
BUFFER
CONTROL PROCESSES
•Stimulus analyzer programs
•Alter biases of sensory channels
•Activate rehearsal mechanism
•Modify information flow from SR to STS
•Code and transfer information from STS to LTS
•Initiate or modify search of LTS
•Heuristic operations on stored information
•Set decision criteria
•Initiate response generator
RESPONSE
GENERATOR
S
E
N R
S E
O G
R I
Y S
T
E
R
STIMULUS
INPUT
RESPONSE
OUTPUT
REHEARSAL
BUFFER
SHORT-TERM
STORE
Memory bank subject
to rapid decay
LONG-TERM
STORE
Self-addressable memory bank
not subject to decay
REHEARSAL
BUFFER
RESPONSE
GENERATOR
S
E
N R
S E
O G
R I
Y S
T
E
R
STIMULUS
INPUT
RESPONSE
OUTPUT
REHEARSAL
BUFFER
SHORT-TERM
STORE
Memory bank subject
to rapid decay
LONG-TERM
STORE
Self-addressable memory bank
not subject to decay
REHEARSAL
BUFFER
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624
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52 x 12
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Figure 6A
Short-term Memory and the Multi-store Memory Model
CONTROL PROCESSES
•Stimulus analyzer programs
•Alter biases of sensory channels
•Activate rehearsal mechanism
•Modify information flow from SR to STS
•Code and transfer information from STS to LTS
•Initiate or modify search of LTS
•Heuristic operations on stored information
•Set decision criteria
•Initiate response generator
RESPONSE
GENERATOR
S
E
N R
S E
O G
R I
Y S
T
E
R
STIMULUS
INPUT
RESPONSE
OUTPUT
REHEARSAL
BUFFER
SHORT-TERM
STORE
Memory bank subject
to rapid decay
LONG-TERM
STORE
Self-addressable memory bank
not subject to decay
REHEARSAL
BUFFER
CONTROL PROCESSES
•Stimulus analyzer programs
•Alter biases of sensory channels
•Activate rehearsal mechanism
•Modify information flow from SR to STS
•Code and transfer information from STS to LTS
•Initiate or modify search of LTS
•Heuristic operations on stored information
•Set decision criteria
•Initiate response generator
RESPONSE
GENERATOR
S
E
N R
S E
O G
R I
Y S
T
E
R
STIMULUS
INPUT
RESPONSE
OUTPUT
REHEARSAL
BUFFER
SHORT-TERM
STORE
Memory bank subject
to rapid decay
LONG-TERM
STORE
Self-addressable memory bank
not subject to decay
REHEARSAL
BUFFER
RESPONSE
GENERATOR
S
E
N R
S E
O G
R I
Y S
T
E
R
STIMULUS
INPUT
RESPONSE
OUTPUT
REHEARSAL
BUFFER
SHORT-TERM
STORE
Memory bank subject
to rapid decay
LONG-TERM
STORE
Self-addressable memory bank
not subject to decay
REHEARSAL
BUFFER
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Figure 6B
Central
Executive
Visuospatial
Sketchpad
Episodic
Buffer
Phonological
Loop
Visual Episodic Language
Semantics LTM
Fluid
Systems
Crystallized
Systems
Baddeley’s Working Memory Module
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Figure 7
Prefrontal CortexWorking Memory
Ventrolateral PFCNon-spatial memory
(color, shape, etc)
Dorsolateral PFCSpatial memory
Anatomical Subdomains of
Working Memory
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Figure 8
Short-term MemoryWorking Memory
Long-term Memory
Action Potential Firing
Sustained Neural Circuit
Activity
Persisting Molecular and
Cellular Changes
Changes in Synaptic
Structure
Anatomical Circuit Alterations
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HUMAN MEMORY
DECLARATIVE
(EXPLICIT)
NONDECLARATIVE
(IMPLICIT)
FACTS EVENTS
PROCEDURAL
(SKILLS AND
HABITS)
PRIMING SIMPLE
CLASSICAL
CONDITIONING
NONASSOCIATIVE
LEARNING
EMOTIONAL
RESPONSES
SKELETAL
MUSCULATURE
MEDIAL
TEMPORAL
LOBE
STRIATUM NEOCORTEX AMYGDALA CEREBELLUM REFLEX
PATHWAYS
Subdivisions of Human Memory
Milner B, Squire LR, Kandel ER: "Cognitive neuroscience and the study of memory". Neuron 1998, 20:445-468.
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• Cognitive Processing:
Space
Time
Relationships
• Memory Consolidation
Functions of the Hippocampus
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From JH Martin, Neuroanatomy
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From JH Martin, Neuroanatomy
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From JH Martin, Neuroanatomy
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Lined Seahorse, Hippocampus erectus
Vertebrate Field Zoology Photo #148, Courtesy Samford University
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From JH Martin, Neuroanatomy
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From JH Martin, Neuroanatomy
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From JH Martin, Neuroanatomy
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Visual Somatosensory
Olfactory
Auditory
Perirhinal Cortex
Entorhinal
Cortex
Dentate Gyrus
CA3
CA1
Sensory Regions
Of Cortex
Perirhinal/
Entorhinal
Cortex
Hippocampal
Formation
Hippocampal Connectivity in the CNS
Parahipp.
Cortex
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Subiculum/
Entorhinal Cortex
Figure 5
Entorhinal
Cortex
Dentate
Gyrus
CA3
CA1
Perforant
Pathway
Mossy
Fibers
Lateral
Septum
Amygdala,
Cortex
Schaffer
Collaterals
Norepinephrine
Acetylcholine
Serotonin
GABAergic Interneuron
CA1 Axon
Schaffer Collaterals
CA3
CA1
Entorhinal Cortex
CA1 Outputs
A B
C
Hippocampal Output Pathway & Intrinsic Circuit
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Figure 4
Hippocampal Pyramidal Neuron
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Major Neurotransmitters in the Hippocampus
Neurotransmitter Abbreviation, Synonym General role
Serotonin 5HT, 5-hydroxytryptamine neuromodulation
Dopamine DA neuromodulation
Acetylcholine ACh neuromodulation
Norepinephrine NE neuromodulation
Gamma-amino GABA inhibitory transmission
Butyric Acid
Glutamate Glu excitatory transmission
Table 4
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• Cognitive Processing:
Space
Time
Relationships
• Memory Consolidation
Functions of the Hippocampus
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Figure 6
H.M. Control
MRI scan of H.M. and control brain
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Surgeon’s Estimate
of H.M.’s lesions
Revised Estimate
based on MRI
MRI of HM’s Brain lesions
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From Eichenbaum and Cohen
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From Eichenbaum and Cohen
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From Eichenbaum and Cohen
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• A makes B happen • Measure
• Activating B should cause C• Mimic
• Blocking B should block A causing C• Block
• None (A makes C happen)• Determine
PredictionExperiment
Hypothesis: A→B→C
Four Types of Experiments
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Virtual Maze
Burgess N, Maguire EA, O'Keefe J.
The human hippocampus and spatial and episodic memory.
Neuron. 2002 35:625-41.
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MRI of Hippocampus
Burgess N, Maguire EA, O'Keefe J.
The human hippocampus and spatial and episodic memory.
Neuron. 2002 35:625-41.
fMRI in
Virtual Maze
Navigation
Morphometry
of London Taxi
Drivers
Right Posterior
Hippocampal
Volume Correlates
with Time
Spent Driving
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Blue Box 1
Neuroimaging of the retrieval of the Spatial Context of an Event
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HUMAN MEMORY
DECLARATIVE
(EXPLICIT)
NONDECLARATIVE
(IMPLICIT)
FACTS EVENTS
PROCEDURAL
(SKILLS AND
HABITS)
PRIMING SIMPLE
CLASSICAL
CONDITIONING
NONASSOCIATIVE
LEARNING
EMOTIONAL
RESPONSES
SKELETAL
MUSCULATURE
MEDIAL
TEMPORAL
LOBE
STRIATUM NEOCORTEX AMYGDALA CEREBELLUM REFLEX
PATHWAYS
Subdivisions of Human Memory
Milner B, Squire LR, Kandel ER: "Cognitive neuroscience and the study of memory". Neuron 1998, 20:445-468.
![Page 45: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/45.jpg)
Figure 10
Human Excellence in Motor Learning and Memory
![Page 46: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/46.jpg)
Medium Spiny Neuron
![Page 47: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/47.jpg)
Figure 11
Summary Scheme of Interaction Between Motor Centers
![Page 48: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/48.jpg)
Figure 12
Basal Ganglia in the Human Brain
![Page 49: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/49.jpg)
Figure 13
Cortex
Striatum
Caudate/Putamen/
Accumbens
Substantia
Nigra
Globus
Pallidus
Subthalamic
Nucleus
Thalamus
Ventral Anterior & Ventral Lateral
Nuclei
Motor Pre-MotorPre-Frontal
(Sensory)
The Striatal Motor Learning System
![Page 50: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/50.jpg)
From Eichenbaum and Cohen
![Page 51: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/51.jpg)
Memory-associated Cognitive Dysfunction in Humans
Disease or Syndrome Type of Memory Affected Likely Anatomical Locus
or Involved*
Schizophrenia Working Memory Prefrontal Cortex
Major Depression Declarative Memory ?
Aging-related Dementias Long-term Declarative and Temporal Lobe Hippocampal
Episodic Memory System
Korsakoff’s Syndrome Long-term Declarative and Mamillary Bodies, Limbic
(Alcoholism, Malnutrition) Episodic Memory System, Thalamus
Huntington’s Disease and Motor Learning, Habit Learning Nigro-striatal System
Parkinson’s Disease
Attention Deficit Generalized or Specific Learning Frontal Lobes?
Hyperactivity Disorder (ADHD) Disabilities, Dyslexia Basal Ganglia?
Post-traumatic Aversive Associative Conditioning, Amygdala, Hippocampus
Stress Disorder* (PTSD) Lack of Extinction
Table 3
![Page 52: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/52.jpg)
Human Excellence in Declarative Learning and Memory
![Page 53: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/53.jpg)
Figure 14
Rajan Srinavasen Mahadevan
![Page 54: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/54.jpg)
3.14159265358979323846264338327950288419716939937510582097494459
2307816406286208998628034825342117067982148086513282306647093844
6095505822317253594081284811174502841027019385211055596446229489
549303819644288109756659334461284756482337867831652712019091456
485669234603486104543266482133936072602491412737245870066063155
8817488152092096282925409171536436789259036001133053054882046652
1384146951941511609433057270365759591953092186117381932611793105
1185480744623799627495673518857527248912279381830119491298336733
624406566430860213949463952247371907021798609437027705392171762
931767523846748184676694051320005681271452635608277857713427577
896091736371787214684409012249534301465495853710507922796892589
2354201995611212902196086403441815981362977477130996051870721134
999999837297804995105973173281609631859502445945534690830264252
2308253344685035261931188171010003137838752886587533208381420617
1776691473035982534904287554687311595628638823537875937519577818
577805321712268066130019278766111959092164201989….
Figure 15
1000 Decimal places of pi
![Page 55: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/55.jpg)
Figure 16
Kim Peek
![Page 56: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/56.jpg)
Figure 2
The Penfield Stimulation Experiments
![Page 57: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/57.jpg)
![Page 58: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/58.jpg)
The Mimic Octopus
![Page 59: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/59.jpg)
![Page 60: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/60.jpg)
http://news
![Page 61: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/61.jpg)
![Page 62: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/62.jpg)
![Page 63: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/63.jpg)
![Page 64: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/64.jpg)
The Mimic Octopus
![Page 65: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/65.jpg)
![Page 66: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/66.jpg)
From JH Martin, Neuroanatomy
![Page 67: Chapter 2: Studies of Human Learning and Memory · Finding Nemo Andrew Stanton (Animated - Ellen DeGeneris as Dory) 2003 50 First Dates Peter Segal (Adam Sandler) 2004 Retrograde](https://reader033.vdocuments.mx/reader033/viewer/2022041621/5e3efa696ed93454ed5dfcaf/html5/thumbnails/67.jpg)
Lateralization of Hippocampal
Function
Burgess N, Maguire EA, O'Keefe J.
The human hippocampus and spatial and episodic memory.
Neuron. 2002 35:625-41.
Studies of hippocampectomy patients
Distance
Traveled
To Correct
Location
Percent
Correct
Impairment
Ratio
Percent
Correct