CHAPTER 10

Learning and memory

OUTLINE

Non- Associative Learning

Associative Learning
Memory

Quiz #2 on Thursday 11/7: study slides 30-

37 (LTD) & 51-62 (LTP)
 Learning
The act of acquiring new, or modifying and reinforcing, existing knowledge,
behaviors, skills, values, or preferences.

Types of learning:
• Non-associative learning
• Associative learning
• Imprinting
• Observational learning
• Play
• Enculturation
 Learning

• Non-associative learning: involve change in the magnitude of response to

environmental events

A B
A B

A B
 Learning

• Non-associative learning: involve change in the magnitude of response to

environmental events

A B
A B

A B
• Associative learning: involve a connection between 2 elements or events

A
A B
 CHAPTER 9
Learning and memory

OUTLINE

Non- Associative Learning

Associative Learning
Memory
 Non-Associative Learning
Learning involving exposure usually to a single event, and that is presumed not
to reflect learning of a relationship between multiple events.

• Habituation
• Dishabituation
• Sensitization
 Non-Associative Learning
Learning with Aplysia

Invertebrate learning offers a simpler system

• simple nervous system (20,000 neurons)
• neuronal development hard-wired
• identifiable individual neurons
 Non-Associative Learning
Learning with Aplysia
 Non-Associative Learning
Habituation:
A decline in responsiveness to repeated stimulation arising from a central
change in the organism.

Gill-Withdrawal Reflex: stimulating the siphon produces a withdrawal

of the gill and siphon.

With repeated stimulation,

the animal will withdraw less
and less.
 Non-Associative Learning
Habituation:
A decline in responsiveness to repeated stimulation arising from a central
change in the organism.

Acoustic startle reflex:

With repeated loud noises, the participant jumps

less and less.
 Non-Associative Learning
Some Characteristics:

Habituation is stimulus specific. Habituation training on one stimulus does not

generalize to other stimuli unless the test stimuli are perceptually similar to the
first.
 Non-Associative Learning
Habituation shows spontaneous recovery when the stimulus that has undergone
habituation is tested after a rest.

REST
 Non-Associative Learning
Habituation is not motor fatigue or sensory adaptation. The decline in
responsiveness is due to a change in the central processes that intervene between
sensory and motor neurons
 Non-Associative Learning
Dishabituation:

Exposure to a new stimulus often disrupts or removes habituation to a prior

stimulus.
 Non-Associative Learning
Sensitization:

The increase in responsiveness to a stimulus that has not undergone habituation

training thought to arise from a general arousal process.

= an increase in behavior due to exposure to a noxious (painful) stimulus

 Non-Associative Learning
Sensitization:

No shock

4 days after
multiple
shocks
 Non-Associative Learning
Sensitization:
 Non-Associative Learning
Sensitization is different from habituation:

Habituation Sensitization

Occurs due to weak Occurs due to strong/painful

stimulation stimulation

Requires repeated stimulation Can occur with a single

stimulation

Decreases behavior Increases behavior

 CHAPTER 10
Learning and memory

OUTLINE

Non- Associative Learning

Associative Learning
Memory
 Associative Learning
Process by which an association between two stimuli or a behavior and a stimulus
is learned.

A
A B
 Associative Learning

• Classical conditioning:
• Pavlov
• Placing a neutral signal before a reflex
• Focuses on involuntary, automatic behaviors
• Operant conditioning
• Skinner
• Applying reinforcement or punishment after a behavior
• Focuses on strengthening or weakening voluntary behaviors
 Associative Learning
Classical conditioning:

A neutral stimulus is repeatedly presented together with a reflex eliciting stimuli

(=unconditioned stimulus) that elicits a behavioral response (unconditioned
response)

REPEATED MULTIPLE TIMES

 Associative Learning

An unconditioned stimulus (US) is a biologically relevant stimulus

An unconditioned response (UR) is an unlearned reaction to the US

 Associative Learning

After conditioning, the neutral stimulus elicits a specific response

 Associative Learning

A conditioned stimulus (CS) is an initially neutral stimulus that acquires the ability
to signal important biological events

A conditioned response (CR) is a learned reaction to the CS

Now Conditioned
stimulus
 Associative Learning
Example of Classical conditioning:

Pavlov: developed an empirical approach for studying classical conditioning

 Associative Learning
Example of Classical conditioning:
 Associative Learning
Example of Classical conditioning:
 Associative Learning
Example of Classical conditioning:
 Start here for Quiz
Associative Learning
Classical conditioning:
Classical eyeblink conditioning
Associative Learning

US pathway:
• Climbing fibers send
information about the US
from the pons/medulla to
the cerebellum.

Pons and medulla

 Associative Learning

CS pathway:
• The pontine nuclei (PN) receive projections
from auditory, visual, somatasensory, and
association systems.
• The PN give rise to mossy fiber (MF) axons
that carry CS-related information to the
cerebellum and terminate at granule cells
(GR) of the cerebellar cortex.
• Granule cells give rise to parallel fiber (PF)
axons which synapse onto Purkinje cells.
 Associative Learning

CS-US Convergence in the Cerebellum

• Purkinje cells of the cerebellum receive
converging CS and US input.
• Cerebellum indirectly activates the red
nucleus
• The red nucleus sends projections to the
facial nucleus = motor output.
 Associative Learning

Long-term depression (LTD) from parallel

fibers onto Purkinje cells

LTD = ability to depress the efficacy of

glutamate transmission

LTD is induced by coupled activation of

two glutamatergic synaptic input
pathways: parallel fibers and climbing
fibers
 Associative Learning

Glutamate bind to both AMPARs and

mGluR1s on the postsynaptic surface
of Purkinje neurons

Activation of mossy fibers by the tone =

release of glutamate
 Associative Learning
Cerebellar Long term Depression (LTD)

Glutamate

AMPAR/mGluR1
(Purkinje cells)

Increase Ca2+
Increase Na+
Activation of protein kinase C (PKC)
 Stop here for Quiz
Associative Learning
Cerebellar Long term Depression (LTD)

Glutamate

AMPAR/mGluR1
(Purkinje cells)

Increase Ca2+
Increase Na+
Activation of protein kinase C (PKC)

Reduced number of glutamate

receptors on Purkinje cells

Less receptors to activate = reduced

EPSP on Purkinje cells from parallel
fibers
 CHAPTER 10
Learning and memory

OUTLINE

Non- Associative Learning

Associative Learning
Memory
 Memory

Memory is the process by which information is encoded, stored, and retrieved.

 Memory

• Encoding : receiving, processing and combining of received information

• Storage: creation of a permanent record of the encoded information

• Retrieval, recall or recollection: calling back the stored information in response

to some cue for use in a process or activity
 Memory

Types of memory:

1. Sensory memory: holds sensory information for a few seconds or

less after an item is perceived.

a. Iconic memory: visual information.

b. Echoic memory: auditory information.
c. Haptic memory: touch stimuli.
 Memory

Types of memory:

2. Short term (working) memory: allows recall for a period of several

seconds to 1-2 minutes. Its capacity is very limited (7±2 items).
 Memory

Brain mechanisms of short-term memory

Joaquin Fuster recorded the electrical activity of neurons in the prefrontal cortex
of monkeys while they were doing a delayed matching task.

1. The monkey sees how the experimenter places a

bit of food under one of two identical looking cups.
2. A shutter is then lowered for a variable delay
period.
3. After the delay, the shutter opens and the monkey
is allowed to retrieve the food from under the cups.

Successful retrieval requires holding the location of the

food in memory over the delay period
 Memory

Brain mechanisms of short-term memory

Cue
End of
delay

firing of certain neurons in the delay

period shows that the brain has a
mechanism of keeping representations
active without external input
 Memory

Brain mechanisms of short-term memory

Prefrontal cortex lesion in rats:

 Memory

Types of memory:

3. Long term memory: can store much larger quantities of information

for potentially unlimited duration.
 Memory

Types of memory:

• Facts • Personal • Riding a

• Verbal experience bike
experiences
 Memory
Brain mechanisms of long-term memory

Temporal lobes: Medial temporal lobe structures that are critical for long-term
memory include:
• amygdala
• Hippocampus
 Memory

Brain mechanisms of long-term memory

- Two-thirds of medial temporal lobes removed in 1953,
including hippocampus, amygdala, and surrounding
cortex

Henry Gustav Molaison

(H.M.): 1926-2008
 Memory

Brain mechanisms of long-term memory

Henry Gustav Molaison (H.M.): 1926-2008

- Profound anterograde amnesia—able to

retain information for only short periods of
time (inability to create new memories)

- Personality unchanged

- Memory highly dependent on attention

 Start here for Quiz
Memory

Brain mechanisms of long-term memory

Hippocampus: plays an important role in the formation of new memories about
experienced event (episodic memory)
 Memory

Brain mechanisms of long-term memory

Hippocampus and long-term potentiation (LTP)
 Memory

Brain mechanisms of long-term memory

Hippocampus and long-term potentiation (LTP)

The brain stores memory by altering the

strength of connections between neurons
that are simultaneously active

= long-term potentiation (LTP): a long-lasting

enhancement in signal transmission between
two neurons that results from stimulating
them synchronously
 Memory

Brain mechanisms of long-term memory

Long-term potentiation (LTP): a long-lasting enhancement in signal transmission

between two neurons that results from stimulating them synchronously

High frequency Low frequency

stimulation stimulation
 Memory
Brain mechanisms of long-term memory
Hippocampus and long-term potentiation (LTP)

The number of ion channels on the

post-synaptic membrane affects the
strength of the synapse

Can last from minutes to days!

 Memory
Brain mechanisms of long-term memory
LTP depends on NMDA receptors activation
 Memory
Brain mechanisms of long-term memory

What happens to your memory if you increase the number of NMDA receptors?
 Memory
Brain mechanisms of long-term memory

Increasing NMDA signaling enhance learning and memory in mice

Mouse with NMDA receptors

that stay open longer

More Ca2+ influx

More robust LTP

Enhanced memory
 Memory
Brain mechanisms of long-term memory

LTP Associativity:

In a weak input, stimulation often fails to induce LTP

No LTP
 Memory
Brain mechanisms of long-term memory

LTP Associativity:

when the weak input is stimulated simultaneously with high-frequency

stimulation of another independent strong input, the weak input can also
exhibit LTP
1

LTP at Pathway 2 when

paired with strong
stimulation at Pathway 1
 Memory
Brain mechanisms of long-term memory

LTP Cooperativity:

LTP can be induced cooperatively via the weaker stimulation of many pathways to
a synapse: when using high frequency stimulation to induce LTP, a crucial
number of presynaptic fibers must be simultaneously activated — they must
'cooperate' to elicit LTP

Simultaneous activity
 Stop here for Quiz
Memory
Brain mechanisms of long-term memory

LTP Associativity & Cooperativity:

 Memory
Brain mechanisms of long-term memory – OTHER BRAIN REGIONS

Diencephalon: the hippocampus is tightly connected to the thalamus. Disruption

to these connections result in amnesia.

Patient N.A.: lesion of the left

dorsomedial thalamus

• Anterograde amnesia (difficulty

forming new memory)

• Preserved short-term memory and

intelligence
 Memory
Cerebral cortex: plays a key role in memory, attention, perceptual awareness,
thought, language, and consciousness. Previously formed memories are thought
to be stored in the cerebral cortex.

Control
Tg mice (NMDAr in cortex)
 Memory
Striatum: role in procedural learning and memory.

Striatum defect
control
 Memory
Brain mechanisms of long-term memory

Summary of brain structures involved in long-term memory

 Memory
Memory loss in aging and Alzheimer’s disease
 Memory
Memory loss in aging and Alzheimer’s disease
Age-related memory loss and Alzheimer’s disease are distinct
conditions
 Memory
Memory loss in aging and Alzheimer’s disease
Age-related memory loss and Alzheimer’s disease are distinct conditions
 Memory
Memory loss in aging and Alzheimer’s disease
Alzheimer’s disease and LTP
Control
Aβ
Aβ + Aβ inhibitor
QUESTIONS?