Transcript Neurobiology of Behavior and Cognition

Neurobiology of
Learning and Memory
Prof. Stephan Anagnostaras
Lecture 2: Learning Theory
Classical (Pavlovian) conditioning
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.
US - unconditional stimulus - biologically significant
stimulus (food)
UR - unconditional response (salivation)
CS - conditional stimulus - previously neutral
stimulus (bell)
CR - conditional response (salivation)
The UR and CR can be different, but it bears some
relationship to the UR.
Classical (Pavlovian) conditioning
After pairing, how do you know you have a CR?
1) Present the CS alone (without the US)
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.
Basic
Phenomena
•Negatively accelerating
growth curve
• The stronger the US,
the stronger the CR
(same growth rate)
Asymptote
Qu ickTime™ and a TIFF (Un compressed) d ecompressor a re needed to s ee th is picture.
Growth
rate
Basic phenomena
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
Basic phenomena
4. Discrimination
• Train CS+ and CS– that are similar
• Inhibition
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
• Discrimination explained using learning theory
• Extinction explained
Basic phenomena
• Inhibition is a weaker process than excitation
• Spontaneous recovery in extinction
• Disinhibition in extinction
• Excitatory association not lost, it’s only the buildup of
inhibition that suppresses excitation
• Law of parsimony
Power of a theory =
# of things explained
---------------------------# of explanatory principles
Procedure, Process, & Behavior
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)
• Our explanation involves all three
• Must be aware of this distinction -- procedure is
not what is learned by the animal
• Skinner argued only talk about procedure-result
laws (radical behaviorism)
Control procedures
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
• Presentation of CS alone increases CR
Pseudoconditioning
Control: present the CS alone
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.
Several acquisition procedures
Qu ickTime™ and a TIFF (Un compressed) decompresso r are needed to see this p icture.
Forward works best. Interestingly this is a test of
Contiguity Theory
Several acquisition procedures
Delay conditioning is
another term for
forward conditioning.
Trace conditioning is
quite special in terms
of mechanistic models
of animal learning.
QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture.
Higher order conditioning
Second-order conditioning
Phase I
Phase II
Test
CS1-US
CS2-CS1
CS2-->CR
tone-food
light-tone
light
Sensory pre-conditioning
Phase I
Phase II
Test
CS2-CS1
CS1-US
CS2-->CR
light-tone
tone-food
light
Generality of conditioning
Conditioning permeates everything you do
can condition pancreas and most glands,
voluntary and involuntary muscles, and immune system
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) 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
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.
Exp 3: Design the same as #2, except female now in
between paired and unpaired male -- female always
picks paired male
Generality of conditioning
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
Qu i c k T i m e ™ a n d a T I F F (U n c o m p re s s e d ) d e c o m p re s s o r a re n e e d e d t o s e e t h i s p i c t u re .
xWhat is learned?
Emotional Learning
• Little Albert study
Conditioned emotional response (CER)
(Pavlovian fear conditioning)
Estes & Skinner (1941) Conditioned Suppression
Trained to bar-press for food
Paired tone with shock
When tone came on fear suppressed bar-pressing
Suppression became the dominant way to measure CR
What is learned?
Why not just measure fear?
• No attention to evolution. Why do rats stop barpressing? They freeze. Nowadays people just measure
freezing or other defensive CR.
E.g. Fanselow & Bolles 1979: Did fear conditioning with
backward (unpaired group)
• Evolution heavily influences what is learned, and even
what can be learned
What is learned?
S-S vs S-R
Two views on learning
S-S: CS--->US---> R
S-R: CS--->R (US serves to stamp in this association)
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?
S-S vs S-R
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
What is learned?
What causes conditioning?
Contiguity theory: things have to occur together, that is
necessary and sufficient
Challenges:
• Simultaneous conditioning doesn’t work well
• Garcia & Koelling (1966)
Conditioned Taste Aversion (CTA)
Typical CTA Procedure
(taste)
CS
(illness inducing agent)
US
Good conditioning
with CS-US delay
of up to 75 min
-Contiguity not
necessary
CR
(disgust)
Avoidance
UR
(illness)
What is learned?
Is contiguity sufficient?
Kamin (1968): Blocking effect
A= CS
+ = US AB+ = two different CSs with US
Train
AB+
light-tone-shock light
However…
Phase I Phase II
A+
AB+
Test
B alone = good conditioning
Test
B alone = no conditioning!!
US must be SURPRISING. Note that contiguity is the
same in both experiments
What is learned?
Is contiguity sufficient?
Un Blocking effect
A= CS
+ = US AB+ = two different CSs with US
Phase I Phase II
A+
AB++
Big US was SURPRISING.
Test
B alone = conditioning!!
What is learned?
It is also surprising if you don’t get the US:
Conditioned inhibition procedure:
Phase I Phase II
Test
A+
AB–
B = cond inhibitor
US was expected but didn’t occur!
Relationship between cue and
consequence
• Garcia & Koelling (1966)
“Bright Noisy Water Experiment”
• taste associated with illness
• audio/visual stimuli associated with
shock
Garcia & Koelling (1966)
Lithium Chloride
Salty w ater
Light
Noise
Garcia & Koelling (1966)
Lithium Chloride
Shock
Taste
Licks
A/V
Taste
A/V
Garcia & Koelling
Biological constraints on learning
Modern learning theory
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
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
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?
Modern learning theory
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
• 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
Correlated group: A perfectly predicts shock, and X
only half the time predicts shock
Uncorrelated group: A predicts shock half the time
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
valid cue in this situation.
Modern learning theory
Rescorla (1968), Contingency experiment
CS = tone, US = shock
For all groups, P (US|CS) = 0.8 (80% of the time you get
the CS you will get the US also).
Rescorla varied the P(US|no CS) for all groups.
Modern learning theory
Qu ickTime™ and a TIFF (U ncompressed) decompresso r are needed to see this pictu re.
Modern learning theory
Rescorla called this contingency theory:
QuickTime™ and a TIFF (Uncompressed) decompress or are needed to see this picture.
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