Neuroscience

10
Fall 2017
Lecture 14: Learning & Memory

MW 4-5:50
Franz 1260
Difference Between Learning and Memory

Learning: The process by which new information is acquired

by the nervous system and is observable through change in
behavior

Memory: The encoding, storage, and retrieval of learned

information
Two Major Classes of Long-Term Memory and
Associated Brain Regions
Long-Term
Memory

Declarative Nondeclarative
(Explicit) (Implicit)

Procedural Associative Non-

Facts Events Priming (Habits/ (Condition- Associative
(Habituation/
Skills) ing) Sensitization)

Emotional Skeletal
Medial
Responses Musculature
Temporal Lobe

Striatum Amygdala Cerebellum Reflex

Neocortex
Pathways

It was only in the 1980s that the idea that not all memories and
memory systems were the same became widely accepted
Differentiating Declarative from Nondeclarative
Memory

Declarative Memory: The storage and retrieval of material

that is available to consciousness and can, in principle, be
expressed by language (events, facts, words, faces, music)

Nondeclarative Memory: Memories that involve skills and

associations that are, by and large, acquired and retrieved
at an unconscious level

Nondeclarative memory is unconscious, task specific, and cannot

be all that flexibly applied, while declarative memory is conscious
and is notable for the flexibility of its application
To Remember or Not Remember?
Whether or not something that is perceived will be
remembered later is determined by a number of factors

1. The number of times the event or fact is repeated

2. The importance of the event or fact
3. Its relationship with knowledge that a person already has
4. The amount of rehearsal of the material after it has first been
presented
 To Remember or Not Remember?
An Example: Chess
Orientation of chess pieces after whites 21st move in game 10 of the 1985 World Chess Championship
After briefly viewing the board from
a real game of chess, master players
can rapidly reconstruct much of the
piece positioning.

A random arrangement of the same 28 pieces

When the same pieces were
randomly configured on a game
board, master players did not
perform any better than novices at
recollecting piece positions.
Conclusion: Retention of briefly
presented information depends on
past experience, context, and
perceived importance
Declarative Memory
Types of Declarative Memory Can Be Divided By
Duration

1. An object or fact can be represented initially in immediate

memory
2. Representation of that object or fact can be sustained in
working memory
3. In certain cases, working memories can be persist as long-
term memory.
Immediate Memory
Immediate Memory: That which can be held actively in
mind beginning the moment that information is received

It is the information that forms the focus of immediate

attention and that occupies the current stream of thought

Capacity limited: Can hold 4-7 things or chunks in

immediate memory

Extremely time limited: Contents persist for <30s unless

actively rehearsed this active rehearsal is called working
memory
Working Memory
Thought to consist of a relatively large set of temporary
capacities each a property of one of the brains
specialized information processing systems
Examples of working memory
Phonological loop: Concerned with language, it temporarily
stores spoken words and meaningful sounds
Supports, for example, the ability to hold in mind a telephone number
while preparing to dial it or the ability to hold in mind words while
speaking or comprehending and ordinary sentence
Visuospatial sketch pad: Stores visual images such as faces and
spatial layouts

There is no single temporary memory store though which

all information moves on its way to long-term memory
Working Memory Relies on the Interplay of Activity
Between the Frontal and Sensory Cortices
The frontal cortex and sensory cortices work together as
a neuronal system to (1) perceive information and then
(2) hold it in working memory
1. Neuronal activity in areas of the higher sensory cortices signals
sensory information that is being perceived at an particular
moment
2. Top-down feedback from the frontal cortex then sustains
neuronal activity in the sensory areas across a delay, biasing
the sensory areas toward stimuli that are important for
ongoing behavior and that need to be held in working memory
Executive Function of Prefrontal Cortex Controls
Access to Working Memory
Executive Function: The ability to direct ones actions
towards future goals
Mediated by dorsolateral and ventrolateral prefrontal cortex

Executive Function
1. Permits memory to be accessed strategically
2. Orchestrates the use of behavior-guiding rules that allow
knowledge relevant to current goals to be brought to mind and
put to flexible use

Absence of executive function leaves individuals

stimulus-bound and able to react only to the immediate
sensory environment
Amnesia
Amnesia is the loss of declarative forms of long-term
memory
Amnesia can be retrograde or anterograde
Retrograde: Memories of the past are lost
Anterograde: The ability to form new memories is lost
Amnesia can result from
Damage to the medial temporal lobe usually results in
anterograde and some retrograde amnesia
Psychological distress (psychogenic) usually results in only
retrograde amnesia for memories about certain topics or
periods of the past
Amnesic patients can
Learn and retain as well as normal subjects in learning motor
skills
Do not differ in attributes such as IQ, motivation, or abstract
thinking, or reasoning
Medial Temporal Lobe (MTL) Structures Involved in
the Formation of Long-Term Declarative Memories
Frontal Parietal Occipital
Cortex Cortex Cortex
Bilateral damage to MTL structures,
even just the hippocampus, impairs
declarative memory
Perirhinal Parahippocampal The amygdala, although a part of the
Cortex Cortex MTL, DOES NOT contribute to the
formation of declarative memories
Entorhinal The cortex of the MTL surrounding the
Cortex hippocampus and amygdala is
important for declarative memory

Hippocampus
Different structures of the medial
temporal lobe likely carry out different
functions
As damage to MTL increases,
Subiculum fewer strategies remain by which
memory can be stored
Medial Temporal Lobe (MTL)
Patient H.M.

In 1953, Henry Molaison (H.M.) had

the much of his hippocampal
formation removed from both medial
temporal lobes in the brain
Surgery resulted in some retrograde
and total anterograde amnesia
without obvious deficits to IQ,
perception, abstract thinking,
reasoning, or motivation
H.M.s ability to learn to trace a star
while viewing his hand and the star
only in a mirror was a watershed
moment demonstrating the memory is
not one thing
Where is the Memory Stored?
Not the Medial Temporal Lobe

While the medial temporal lobe is essential for forming new

declarative memories, patients with damage to this region
experience temporally graded retrograde amnesia where
more distant memories of the past depend less upon the
medial temporal lobe being intact
Where is the Memory Stored?
The Cortical Engram
While the medial temporal lobe is essential for forming
new declarative memories, patients with damage to this
region experience temporally graded retrograde amnesia
where more distant memories of the past depend less
upon the medial temporal lobe being intact

Perception of an object requires the simultaneous

activation of neurons in many different regions of cortex
Long-term memories of these perceptions are likely
stored in the same distributed assembly of cortical
structures that are engaged in the initial perception and
processing of what is to be remembered
Engram: The sum total of changes in the brain that first
encoded an experience and then constitute the record of
an experience
Where is the Memory Stored?
Lashleys Mass Action Principle
Lashleys Mass Action Principle: Degradation in learning
and memory depends on the the amount of cortex
removed and the more complex a task is, the more
disruptive a lesion of given size is
Memory Retrieval/Forgetting
Retrieval
Recall, i.e., remembering, depends upon the quality of recall cue that triggers
remembering as well as the strength of the original memory
The most effective cues are the ones trigger the best-encoded aspects of the
event being remembered
Forgetting
Ordinary Forgetting: The normal loss of memory strength over time
Retention of memories can be influenced by rehearsal or intentional non-
rehearsal
Individuals with hyperretensive memory can be adversely effecting
In animals, the cellular substrates of memories, e.g., synaptic strength can be
seen to degrade over time
Most likely, previously existing memories are affected by (1) the new storage of
information continually resculpting previously existing memories and (2) the
passage of time itself.
Clive Wearing Composer Devastated by Herpes
Encephalitis
https://www.youtube.com/watch?v=LlNB7dAXQEc
Marilu Henner 60 Minutes
https://www.youtube.com/watch?v=6Dik5YFwoRs
Nondeclarative Memory
Two Major Classes of Long-Term Memory and
Associated Brain Regions

Long-Term
Memory

Declarative Nondeclarative
(Explicit) (Implicit)

Procedural Associative Non-

Facts Events Priming (Habits/ (Condition- Associative
(Habituation/
Skills) ing) Sensitization)

Emotional Skeletal
Medial
Responses Musculature
Temporal Lobe

Striatum Amygdala Cerebellum Reflex

Neocortex
Pathways
Priming: An improvement in the ability to identify or
process words or objects after recent experience with them
A Typical Priming Test
Time 1 Time 2 Completion Rate
Read Test
70

Stems completed with target words (%)

List A

element ter _______ Priming

corduroy pen ______ 50
pleasant cor ________
technical tec ________
cro ________ 30
ele ________
Not read pil ________
List B ple ________ 10

pendant
crocodile List A List B
pillow (primed) (unprimed)
terrain
Priming is Non-declarative
Amnesics could perform as well as control subjects on the
task . . .
Use this three-letter word stem to form the first word that
comes to mind
BUT NOT . . .
Use this three-letter cue to remember a recently
presented word

Even though, in both cases, the study subjects had been

presented a list of words previously that contained these
word stems
Perceptual Learning: Developing an improvement in the
ability to discriminate sensory stimuli, including simple
things like tones or line orientations merely as the result of
performing the discrimination repeatedly.
 Is this group
oriented vertically
or horizontally?
Improved ability to
identify specific
stimulus lasts years!
Observations from Perceptual Learning Experiments

1. Reward or feedback about errors not required

2. When target was moved to the opposite quadrant on
the same trained side as the original training, all
improvements were eliminated
3. When the orientation of the background elements was
changed (from vertical to horizontal or horizontal to
vertical) improvements were lost
4. When initial training was done with only one eye open,
improvements did not transfer to the other eye

Therefore, the likely region(s) involved with perceptual

learning are primary and early secondary sensory cortices!
Emotional Learning: Attaching a feeling of emotion, positive
or negative, when evaluating a piece of information is to a
large extent and unconscious (nondeclarative) product of
learning
A Example of Emotional Learning: Auditory Tone-
Foot Shock Pairing
Lesions of the amygdala interfere with the conditioning of fear
responses to both cue (tone) and context (cage), whereas lesions to
declarative memory regions like the hippocampus interfere with
conditioning to the context but not to the cue
The amygdala is thus involved in the conditioning of fear responses
to simple, modality-specific conditioned stimuli as well as to
complex, polymodal stimuli, whereas the declarative memory
regions such as the hippocampus are only involved in fear
conditioning situations involving complex, polymodal events.
Thus, the amygdala plays an associative in fear conditioning, while
declarative memory regions play more of a sensory relay role

Therefore, it is possible to develop adverse emotional associations

with people, places, and things without any explicit memory of the
events that caused these associations in the first place. This is
emotional memory.
A Example of Emotional Learning: Auditory Tone-
Foot Shock Pairing

Information processed by the auditory centers in the rodent brainstem is relayed to the
auditory cortex via the medial geniculate complex (1). The amygdala receives auditory
information indirectly via the auditory cortex (2) and directly from one subdivision of
the medial geniculate (3). The amygdala also receives sensory information about other
sensory modalities, including pain (4). Thus, the amygdala is in a position to associate
diverse sensory inputs, leading to new behavioral and autonomic responses to stimuli
that were previously devoid of emotional content (5).
Non-Emotional Associative Conditioning
Associative Conditioning: The generation of a novel
response that is gradually elicited by repeatedly pairing a
novel stimulus with a stimulus that normally elicits the
response being studied

Two Broad Forms of Associative Conditioning

Classical conditioning: When an innate reflex is modified
by associating its normal triggering stimulus with an
unrelated stimulus by repeated association
Operant conditioning: The altered probability of a
behavioral response caused by associating the response
with a reward
Classical Conditioning An Example
Delay Eye-blink Conditioning - Richard Thompson
https://www.youtube.com/watch?v=7yfN9ZHIB2o
The Delay Eye-blink Conditioning Circuit Place of
Association

Place of
Tone is the conditioned stimulus (CS) Engram
Air puff is unconditioned stimulus (US) Storage
CS-US association is made at the Purkinje neuron, where
Multiple pairings of CS and US are used to the tone (CS) is carried by the mossy-fiber/parallel-fiber
train the eye-blink response (Delay Block) input and the air puff (US) is carried by the climbing fiber
input from the inferior olive
Engram of memory stored in interpositus deep cerebellar
nucleus
Procedural Learning
When Milner showed that H.M. could
learn to trace star using only the
reflection of his hand in a mirror,
scientists had some of the first
information that not all memories
were the same or handled by the same
brain regions

Brain areas involved in procedural

learning include motor cortex,
striatum, prefrontal cortex, cerebellum
Habituation/Sensitization

Habituation: The animal becomes less responsive to

repeated occurrences of the same stimulus
Human example: Ignoring the feel of clothes

Sensitization: The animal generalizes an aversive response

elicited by a noxious stimulus to a variety of of other non-
noxious stimuli
Human example: Increased motor responses to repeated
presentation of stimulants like cocaine or amphetamines