Chapter 2:

Studies of Human

Learning and Memory

From Mechanisms of Memory, second edition

By J. David Sweatt, Ph.D.

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

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

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.

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

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

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

624

52 x 12

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

Figure 6B

Central

Executive

Visuospatial

Sketchpad

Episodic

Buffer

Phonological

Loop

Visual Episodic Language

Semantics LTM

Fluid

Systems

Crystallized

Systems

Baddeley’s Working Memory Module

Figure 7

Prefrontal CortexWorking Memory

Ventrolateral PFCNon-spatial memory

(color, shape, etc)

Dorsolateral PFCSpatial memory

Anatomical Subdomains of

Working Memory

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

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.

• Cognitive Processing:

Space

Time

Relationships

• Memory Consolidation

Functions of the Hippocampus

From JH Martin, Neuroanatomy

From JH Martin, Neuroanatomy

From JH Martin, Neuroanatomy

Lined Seahorse, Hippocampus erectus

Vertebrate Field Zoology Photo #148, Courtesy Samford University

From JH Martin, Neuroanatomy

From JH Martin, Neuroanatomy

From JH Martin, Neuroanatomy

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

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

Figure 4

Hippocampal Pyramidal Neuron

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

• Cognitive Processing:

Space

Time

Relationships

• Memory Consolidation

Functions of the Hippocampus

Figure 6

H.M. Control

MRI scan of H.M. and control brain

Surgeon’s Estimate

of H.M.’s lesions

Revised Estimate

based on MRI

MRI of HM’s Brain lesions

From Eichenbaum and Cohen

From Eichenbaum and Cohen

From Eichenbaum and Cohen

• 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

Virtual Maze

Burgess N, Maguire EA, O'Keefe J.

The human hippocampus and spatial and episodic memory.

Neuron. 2002 35:625-41.

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

Blue Box 1

Neuroimaging of the retrieval of the Spatial Context of an Event

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.

Figure 10

Human Excellence in Motor Learning and Memory

Medium Spiny Neuron

Figure 11

Summary Scheme of Interaction Between Motor Centers

Figure 12

Basal Ganglia in the Human Brain

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

From Eichenbaum and Cohen

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

Human Excellence in Declarative Learning and Memory

Figure 14

Rajan Srinavasen Mahadevan

3.14159265358979323846264338327950288419716939937510582097494459

2307816406286208998628034825342117067982148086513282306647093844

6095505822317253594081284811174502841027019385211055596446229489

549303819644288109756659334461284756482337867831652712019091456

485669234603486104543266482133936072602491412737245870066063155

8817488152092096282925409171536436789259036001133053054882046652

1384146951941511609433057270365759591953092186117381932611793105

1185480744623799627495673518857527248912279381830119491298336733

624406566430860213949463952247371907021798609437027705392171762

931767523846748184676694051320005681271452635608277857713427577

896091736371787214684409012249534301465495853710507922796892589

2354201995611212902196086403441815981362977477130996051870721134

999999837297804995105973173281609631859502445945534690830264252

2308253344685035261931188171010003137838752886587533208381420617

1776691473035982534904287554687311595628638823537875937519577818

577805321712268066130019278766111959092164201989….

Figure 15

1000 Decimal places of pi

Figure 16

Kim Peek

Figure 2

The Penfield Stimulation Experiments

The Mimic Octopus

http://news

The Mimic Octopus

From JH Martin, Neuroanatomy

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