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What are animals really learning? The case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational and neural perspectives Case in point: Reinstatement Acquisition Extinction no shock Test no shock no shock What can they be learning?

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Page 1: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

What are animals really learning? The case of extinction

PSY/NEU338: Animal learning and decision making: Psychological, computational and neural perspectives

Case in point: Reinstatement

Acquisition Extinction

no shock

Test

no shock

no shock

What can they be learning?

Page 2: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

Extinction ≠ Unlearning

Test

Storsve, McNally & Richardson, 2012

Acquisition Extinction

no shock

Extinction ≠ Unlearning

Acquisition(Context A)

Extinction(Context B)

no shock

Test (renewal)

Bouton & Bolles 1979

free

zing

(s

uppr

essi

on r

atio

)

Page 3: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

And… this depends on integrity of hippocampus

Ji & Maren 2005

Extinction does something, but not what we want it to do

Page 4: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

Extinction does something, but not what we want it to do

Redish: Animals also learn the states of the task

idea: persistently negative prediction errors (low reward rate) cause splitting

of new state

Page 5: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

But: this does not explain similar phenomena in LI

no LI if conditioning is in different context than

preexposure

Honey & Good 1993

and this too depends on hippocampus… What is

the hippocampus doing in these tasks?

going one step further: learning causal structure

structure I: 

tone causes shock 

structure II: 

latent variable (y) 

causes tone and shock 

structure III: 

tone and shock caused 

by independent latent 

variables (y,z) 

Sam Gershman

Page 6: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

Bayesian inference with an infinite capacity prior

Hypothesis: Animals assume a generative model that is flexible enough to capture the complex structure of the environment, but constrained enough to allow learning

1. Each trial is caused by a single latent cause

2. Each latent cause tends to produce similar trials (ie, has a characteristic probability of “emitting” each of the stimuli)

3. All else equal, a (recently) prolific latent cause (ie, has caused many trials) is more likely to cause another trial

4. The number of possible latent causes is unbounded. That is, there is some (small) probability that the current trial is generated by a completely new latent cause

Gershman, Blei & Niv 2010

Chinese Restaurant Prior (Infinite capacity mixture model)

Page 7: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

Acquisition(Context A)

Extinction(Context B)

Test(Context A)

x y

1. latent causes have characteristic emission probabilities2. similarity between trials allows inference about the relevant

latent cause

A latent cause theory

Inference: “inverting” a generative model

...

generative process

Page 8: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

infe

renc

e (B

ayes

rul

e)

...

p(cause|data) ∝ p(data|cause)p(cause)

Inference: “inverting” a generative model

Bouton & Bolles 1979

Model explains renewal of fearGershman, Blei & Niv 2010

Page 9: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

Conditioning as clustering

Within each cluster: “learning as usual”

(Rescorla-Wagner, RL etc.)

Equivalent to multiple associations

w

reinforcement learning (RW/TD) model

latent-cause modulated RL model

w2

w1

latent cause

Page 10: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

Equivalent to “compound cues”

w

reinforcement learning (RW/TD) model

latent-cause modulated RL model

C1 w

C2 w

Latent causes as states

w

reinforcement learning (RW/TD) model

latent-cause modulated RL model

C1 w

C2 w

Page 11: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

similarity is crucial for clustering observations

inference about latent causes determines “internal state”

Testing the model I : “Circles Task”

how many circles?

Gershman & Niv 2013

Page 12: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

0 50 1000

0.02

0.04

..

number of circles p

rob

ab

ility

condition 1

0 50 1000

0.02

0.04

.

.

number of circles

pro

ba

bili

ty

condition 2

Testing the model I : “Circles Task”

60

61

62

63

64

65

mea

n gu

ess

(blu

e tr

ials

)

cond 1 cond 2

p<0.05

Gershman & Niv 2013

A “battle” between learning and memory

associative learning (modify existing cluster)

Page 13: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

structural learning (create new state)

A “battle” between learning and memory

Testing the model II: how to erase a fear memory

hypothesis: prediction errors (dissimilar data) lead to new states

acquisition extinction

what if we make extinction a bit more similar to acquisition?

Page 14: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

change (prediction error)

wei

ght

chan

ge

(ori

gina

l mem

ory)

new

late

nt c

ause

infe

rred

Testing the model II: how to erase a fear memory

regular extinction

Testing the model II: gradual extinction

acquisition

gradual extinction

gradual reverse

extinction

test one day (reinstatement) or 30 days later (spontaneous recovery)

Gershman, Jones, Norman, Monfils & Niv 2013

Page 15: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

Testing the model II: gradual extinction

first trials of extinction

Pre−cs 1 2 3 40

0.2

0.4

0.6

0.8

1Fr

eezi

ngEXT start

StandardGradualGradual reverse

last trials of extinction

1 2 3 40

0.2

0.4

0.6

0.8

1

Free

zing

EXT end

StandardGradualGradual reverse

Gershman, Jones, Norman, Monfils & Niv 2013

Testing the model II: gradual extinction

only gradual extinction group shows no reinstatement

Gershman, Jones, Norman, Monfils & Niv 2013

Page 16: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

Testing the model II: gradual extinction

only gradual extinction group freeze LESS after 30 days than at the end of extinction (= no spontaneous recovery)

Gershman, Jones, Norman, Monfils & Niv 2013

Where does this “clustering” occur?

key decision: is current observation (trial) similar or different from previous observations?

an expert in such decisions: HIPPOCAMPUS

Page 17: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

Where does this “clustering” occur?

key decision: is current observation (trial) similar or different from previous observations?

an expert in such decisions: HIPPOCAMPUS

Ji & Maren 2005

Where does this “clustering” occur?

Yap & Richardson 2007

acquisition/extinction/test

Page 18: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

But now the really crazy stuff: Monfils-Schiller paradigm

But now the really crazy stuff: Monfils-Schiller paradigm

Page 19: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

In case you are skeptic: this also works in humans...

Even a year later!

In case you are skeptic: this also works in humans...

Page 20: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

An explanation: reconsolidation

• Animals might be smarter than we think: make inferences about the latent causes of observations as part of prediction learning

• similarity determines when different observations will interfere with each other: create a new memory or alter an old one? Inferred structure determines the “battle” between learning and memory in the brain

• we can control memory modification by titrating prediction errors

• memory is not just a passive record: depends on the animal’s beliefs about the underlying causal structure (subjective!)

• potential interaction between dopamine and hippocampus in reinforcement learning

How many circles? 

Summary

Page 21: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

Bayesian inference: what is all the hype about?

• random variables (discrete, continuous)

• probability distribution

• probability distributions as beliefs

• Bayes rule

Bayesian inference: intuition

• sometimes P(A|B) is really hard but P(B|A) is easy → this is why inversion using Bayes rule helps

• eg. you saw 3 heads and 7 tails, what is the probability that the coin is fair?

• If the coin is fair, what is the probability of seeing 3 heads and 7 tails?

• eg. Brian says: “I’ll pass” - what is Brian talking about?

• If Brian is talking about an exam, how probable is this sentence?

P(A|B) = P(B|A)P(A)/P(B)P(model|data) = P(data|model)p(model)/p(data)

Page 22: What are animals really learning? The case of extinctionyael/PSY338/18 Extinction online.pdfThe case of extinction PSY/NEU338: Animal learning and decision making: Psychological, computational

The importance of priors in Bayesian inference

• interpretation of Bayes rule: we care about both prior and likelihood in inference

• eg. test for disease came out positiveaccuracy of test is 99%disease a-priori in 1/10000 people what are the odds that the patient has the disease?