Download Neurobiology of Learning and Memory

Prof. Stephan Anagnostaras
Lecture 2: Learning Theory
Classical (Pavlovian) conditioning
Classical (Pavlovian) conditioning
Neurobiology of
Learning and Memory
Twitmyer (1902)
Paired bell with patellar tendon tap
• Previously neutral bell could now elicit knee jerk
Ivan Pavlov
Studied digestion, and noticed that after he worked
with a particular dog for a while, the dog salivated
when it first saw him.
Paired metronome with food
• Previously neutral metronome elicited salivation.
• Called this conditioning
Classical (Pavlovian) conditioning
A conditional relationship emerged between the
meaningful and previously neutral stimulus.
After pairing, how do you know you have a CR?
1) Present the CS alone (without the US)
US - unconditional stimulus - biologically significant
stimulus (food)
UR - unconditional response (salivation)
CS - conditional stimulus - previously neutral
stimulus (bell)
CR - conditional response (salivation)
2) Measure the response at the beginning of the CS
(metronome) before the US is presented (food)
• One theory is that the purpose of CSs is to predict
USs and the CR is a prepatory response.
The UR and CR can be different, but it bears some
relationship to the UR.
Basic phenomena
Basic
Phenomena
Asymptote
•Negatively accelerating
growth curve
• The stronger the US,
the stronger the CR
(same growth rate)
Growth
rate
1.Acquisition from CS–US pairings
• the curve is negatively accelerating
• the stronger US produces a higher asymptote
• the CR gets stronger with repeated trials
2. Extinction
• the CS is presented alone after conditioning• CS–
• same curve as acquisition
• not unlearning or erasing memory
3. Generalization
• if you present a similar CS you will get a similar
reaction
• generalization decrement
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Basic phenomena
Basic phenomena
4. Discrimination
• Train CS+ and CS– that are similar
• Inhibition
• Inhibition is a weaker process than excitation
• Spontaneous recovery in extinction
• Disinhibition in extinction
Associative learning theory
• Tries to explain what is going on and relies on 3
processes to explain everything
1. Excitation (excitatory association)
2. Inhibition (inhibitory association)
3. Generalization
• Excitatory association not lost, it’s only the buildup of
inhibition that suppresses excitation
Procedure, Process, & Behavior
# of things explained
---------------------------# of explanatory principles
Control procedures
• Discrimination explained using learning theory
• Extinction explained
• Law of parsimony
Power of a theory =
Procedure = what we do (e.g., pair CS and US)
Process = what intervenes between procedure and
behavior (e.g., excitation, inhibition)
Behavioral result what we observe (e.g., after
extinction we see a reduction of the CR)
In order to study associative learning, must show
change in behavior is due to pairing of the CS and
US
• Presentation of stimulus alone increases CR:
Sensitization
Control: present the US alone
• Our explanation involves all three
• Must be aware of this distinction -- procedure is
not what is learned by the animal
• Presentation of CS alone increases CR
Pseudoconditioning
Control: present the CS alone
• Skinner argued only talk about procedure-result
laws (radical behaviorism)
Several acquisition procedures
Control procedures
How could we combine the two control groups?
Unpaired group receives both the US
(sensitization) and CS (pseudoconditioning) but
not together.
Alternative is the truly random control.
The main point is subject has same experience with
CS and US as the Conditioning group.
Forward works best. Interestingly this is a test of
Contiguity Theory
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Several acquisition procedures
Higher order conditioning
Delay conditioning is
another term for
forward conditioning.
Second-order conditioning
Trace conditioning is
quite special in terms
of mechanistic models
of animal learning.
Phase I
Phase II
Test
CS1-US
CS2-CS1
CS2-->CR
tone-food
light-tone
light
Sensory pre-conditioning
Generality of conditioning
Phase I
Phase II
Test
CS2-CS1
CS1-US
CS2-->CR
light-tone
tone-food
light
Generality of conditioning
Coke (CS)-----> Sugar US----> UR (insulin release)
…after a few pairings…
Coke (CS) ---> CR (insuline release)
• Abrupt switch to Diet Coke can cause hypoglycemia
• Pavlovian conditioning prepares the body for
impending URs
Generality of conditioning
Hollis (1989)
Exp 2:
Paired: light (CS) paired with access to females (US)
Unpaired: light unpaired with access to females
Testing: get light then access to female
Result: when light turned on paired group started mating
much more rapidly than unpaired.
Hollis (1989) blue gouramis mating behavior - if a male
enters territory drives it away
Exp 1:
• Males were subjects
• Training:
Paired: light (CS) paired with access to males (US)
Unpaired: light unpaired with access to males
Testing: the light was turned on and barrier removed.
Paired male always won against unpaired male. But also
drives away female.
Generality of conditioning
Conditioning permeates everything you do
can condition pancreas and most glands,
voluntary and involuntary muscles, and immune system
Hollis (1997)
Exp 4: Reproductive success
Training: Paired got light with access to female for
2h, Unpaired got light unpaired with access.
Testing: present light then
give access to female for
2 h for both groups.
Six days later count
baby gouramis
Exp 3: Design the same as #2, except female now in
between paired and unpaired male -- female always
picks paired male
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xWhat is learned?
What is learned?
Emotional Learning
Why not just measure fear?
• Little Albert study
Conditioned emotional response (CER)
(Pavlovian fear conditioning)
• No attention to evolution. Why do rats stop barpressing? They freeze. Nowadays people just measure
freezing or other defensive CR.
Estes & Skinner (1941) Conditioned Suppression
Trained to bar-press for food
Paired tone with shock
When tone came on fear suppressed bar-pressing
E.g. Fanselow & Bolles 1979: Did fear conditioning with
backward (unpaired group)
• Evolution heavily influences what is learned, and even
what can be learned
Suppression became the dominant way to measure CR
What is learned?
What is learned?
S-S vs S-R
S-S vs S-R
Two views on learning
Rescorla (1974) Inflation experiment
1. Tone-shock (0.5 mA)
2. US alone groups:
- 3 mA
- 1 mA
- 0.5 mA
- no shock
3. Test CS alone
- little devaluation in 0.5 mA group
- massive inflation in 1 and 3 mA groups
- Memory of the shock changed and CR changed
Strong evidence for S-S learning:
Rescorla (1973): Devaluation Experiment
•Conditioned Suppression
1. Light (CS) paired with loud noise (US)
2. US alone - habituate (control = no habituation)
3. Test to CS
- habituation group much less fear
What is learned?
What causes conditioning?
Typical CTA Procedure
S-S: CS--->US---> R
S-R: CS--->R (US serves to stamp in this association)
(taste)
(illness inducing agent)
CS
US
Contiguity theory: things have to occur together, that is
necessary and sufficient
-Contiguity not
necessary
Challenges:
• Simultaneous conditioning doesn’t work well
• Garcia & Koelling (1966)
Conditioned Taste Aversion (CTA)
Good conditioning
with CS-US delay
of up to 75 min
CR
(disgust)
UR
(illness)
Avoidance
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What is learned?
What is learned?
Is contiguity sufficient?
Is contiguity sufficient?
Kamin (1968): Blocking effect
A= CS
+ = US AB+ = two different CSs with US
Un Blocking effect
A= CS
+ = US
Train
AB+
light-tone-shock light
However…
Phase I Phase II
A+
AB+
Test
B alone = good conditioning
Test
B alone = no conditioning!!
AB+ = two different CSs with US
Phase I Phase II
A+
AB++
Test
B alone = conditioning!!
Big US was SURPRISING.
US must be SURPRISING. Note that contiguity is the
same in both experiments
What is learned?
Relationship between cue and
consequence
It is also surprising if you don’t get the US:
Conditioned inhibition procedure:
Phase I Phase II
Test
A+
AB–
B = cond inhibitor
• Garcia & Koelling (1966)
“Bright Noisy Water Experiment”
US was expected but didn’t occur!
• taste associated with illness
• audio/visual stimuli associated with
shock
Garcia & Koelling (1966)
Garcia & Koelling (1966)
Lithium Chloride
Lithium Chloride
Salty water
A/V
Taste
Licks
Light
Noise
Shock
Taste
A/V
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Garcia & Koelling
Modern learning theory
Biological constraints on learning
Wagner, Logan, Haberlandt & Price (1968)
Relative validity Theory
Two cmpd CSs: AX (tone, light),BX (buzzer, light)
Animal sometimes get AX, sometimes BX
In group 1 (correlated conditioning group): AX is
reinforced 100% (AX+) and BX is never reinforced (BX-)
In group 2 (the uncorrelated group): AX is reinforced
50% of the time, and BX is reinforced 50% of the time.
Modern learning theory
Modern learning theory
In training…
Correlated group
AX = 100% reinf
BX = 0%
Uncorrelated Grp
AX = 50%
BX = 50%
A predicts US
B predicts no US
neither A or B
perfectly predicts US
In test phase:
Correlated gp
Uncorrelated gp
A alone
Strong cond
No cond
B alone
No Cond
No cond
X alone
No cond
Strong cond
X has the same number of pairings in both groups, so
contiguity theory is screwed
Modern learning theory
• Wagner says the cue must be the most valid predictor
of the US in the situation in order to get associated.
Relative validity to other CSs.
Modern learning theory
Both get 50% reinforcement overall. But what is
happening to X? X is reinforced 50% of the time in both
groups. According to contiguity theory should have the
same conditioning. What happens?
Correlated group: A perfectly predicts shock, and X
Rescorla (1968), Contingency experiment
only half the time predicts shock
CS = tone, US = shock
For all groups, P (US|CS) = 0.8 (80% of the time you get
Uncorrelated group: A predicts shock half the time
the CS you will get the US also).
when its on, the same for B. But X predicts shock half
the time whether A or B are on or not. So X is the most
Rescorla varied the P(US|no CS) for all groups.
valid cue in this situation.
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Modern learning theory
Modern learning theory
Rescorla called this contingency theory:
If P(US|CS) > P (US|no CS) then excitatory conditioning
If P(US|CS) < P(US|no CS) then inhibitory conditioning
(e.g., safety signal)
If P(US|CS) = P(US|no CS) then no conditionin occurs
(truly random control)
Rescorla-Wagner Model (1972)
Key Assumptions:
1. Emphasize CS-US pairings as criticial for
conditioning
2. Formalize the notion of Kamin’s suprirse
3. Assume that any US can only support a limited
amount of conditioning/reinforcement
4. All the CSs compete with echother for the
limited amount of conditioning/reinforcement
5. Competition occurs through summation of all
the CSs present on a given trial
•The US has a certain amount it can condition,
meaning this is a US-limiting model.
•Stimuli compete for ability to predict the US.
Rescorla-Wagner
Model (1972)
Rescorla-Wagner Model (1972)
Can explain a number of phenomena:
Acquisition, extinction
Blocking (A+, AB+, … B)
Unblocking (A+, AB++, …B)
Conditioned Inhibition (A+, AB–, … B)
Contingency
• Can deal with a number of phenomena and makes
several new predictions which were testable
• Cannot deal with latent inhibition (CS pre-exposure)
• Can deal with US pre-exposure effect
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