Classical Conditioning Learning & Memory Arlo Clark-Foos What is classical conditioning?

• Learning to associate previously neutral stimuli with the subsequent events.

• Howard Eichenbaum’s Thanksgiving

Pavlov’s psychic secretion Are you conditioned?

• Some examples of every day conditioning… – Holiday Traditions – Food Associations – Fears – Superstitions – Habits – Skills? Ivan Pavlov

• How are digestive fluids controlled? – Historical view – Pavlov’s view

– Pavlov’s Original Experiment Pavlov’s Experiments

• Psychic Secretion – Specialized procedure for introducing food • Claude Bernard’s psychic secretion in horses – Pavlov’s psychic secretion was unreliable but…

Pavlov Museum, Ryazan, Russia Stimuli and Responses

This is appetitive conditioning. What is an example of aversive conditioning? Conditioned Emotional Response

Estes & Skinner

Appetitive

(Dudai, Jan, Byers, Quinn, & (Domjan, Lyons, North, & Benzer, 1976) Bruell, 1986) Slapping and Blinking in the Name of Research

Ernest Hilgard Clark Hull

Electromyography (EMG) Photo Sensors

Very well studied Rabbit Eyeblink Conditioning

reactive

predictive It gets more complicated…

• Similarity among species • Tolerance, compensatory responses, and homeostasis • Stimulus Timing and Presentation – Contemporaneous Presentation • Not spaced too far apart in time – Is there an ideal spacing? – Order and Consistency • Reliable relationship/expectation Conditioning Procedures

Interstimulus Intertrial

Interval Interval

Forward Conditioning Learning Not to Respond

Conditioned Inhibition: Decrease in CR in response to CS. – Need Baseline

CRs diminish CS+1(Tone)  US CS1 ? CS1 + CS2  over time as CS- CS-2(Light)  CS2 ? inhibits CS+

Baseline Transfer of Learning

• Generalization

CS800mHz  CR CS  Max CR CS(Tone,1200mHz)  US 1200mHz CS1600mHz  CR

• Discrimination

CS(Tone,300mHz)  CS300mHz CS(Tone,500mHz)  US CS500mHz  CR CS(Tone,800mHz)  CS800mHz Context as CS

• Penick & Solomon (1991) – Eyeblink conditioning in rats – Hippocampal Lesions

Transfer Appropriate Processing & Encoding Specificity What is being conditioned?

• How is it learned and what is the nature of the association? S-S or S-R Association?

• Stimulus Substitution Theory (Pavlov) – Definition (S-R Association) – US, CS, and Response centers in the brain

US Response S-S Association

CS S-R Association

– Problem: a CR is not a UR • CR eyeblink is often more gradual and less complete Rescorla (1973)

US Devaluation

Conditioned Suppression Habituate Noise Lever  Reward Lever + Light? (Light/CS + Loud Noise/US)

US

Response S-S Less CR after US devaluation. Association CS S-R S-S Association Association After Conditioning

• After learning, what happens when you present the CS alone?

Extinction What happens in extinction?

• What do we (researchers) see? – No CR = Forgetting?

– Excitatory and Inhibitory Associations (Pavlov) • CC • Extinction

Human eyeblink conditioning and the reduction in responses during extinction. Extinction = Forgetting?

• Spontaneous Recovery – Pavlov: Inhibitory connections are weak, fade – Alt. Theory: Attention/Interest in CS (habituation?)

Human eyeblink conditioning and the reduction in responses during extinction. Extinction = Forgetting?

• Disinhibition – Surprising, typically arousing, new stimulus – Return of CR (akin to sensitization)

• Rapid Reacquisition – Retraining vs. Original Conditioning – Something is retained

Extinction is NOT Forgetting Compound Conditioning

• Context, Multiple Cues – Extinction: “respond” and “don’t respond” • Overshadowing – Salience Error Correction

• Problems with Aristotle’s contiguity

• Informational value of cues

• Kamin’s (1969)

Compound conditioning Rescorla-Wagner (1972)

• Learning on Trains – Contiguity is not enough

• Competition for associative strength

• Prediction Error – Positive vs. Negative prediction errors – Error-correction learning R-W in Humans

• Error Correction in Human Category Learning – Bower & Trabasso, 1964 • Informational value of dot Modelling Conditioning

• Associative Weights – Connectionist Models (e.g., McClelland & Rumelhart) Pay Attention!

• Exposure to CS alone retards later learning – Attention to stimuli – Latent inhibition • Lubow & Moore (1959) – Sheep and Goats • Where is the surprise?

– US Modulation Theory • Prediction error (Rescorla & Wagner, 1972)

– CS Modulation Theory • Attention to stimuli (Mackintosh, 1975) Properties of C Conditioning

• It takes time – 4-5 mo., no eyeblink conditioning in first block but exposure necessary for later learning.

• Other Factors – Intensity of CS-US – Timing, ISI

Ivkovich et al., 1999 Intensity of CS-US

• Faster and More effective

US Intensity Maintained

US Intensity Decreased Timing, ISI

• Timing is critical! – Ideal ISI for rapid learning – Humans = Animals Pavlov observed no CRs with Backward Conditioning: CS does not predict US.

Animals must be learning association AND temporal contiguity Taste Aversion (Garcia Effect)

• Temporal Contiguity – Food poisoning after a date…

• Belongingness: CS-US pairings. – Tone + Food  Shock or Poison – (Garcia & Koelling, 1966)

• Neurological basis: gustatory cortex

• Coyotes (Gustavson et al., 1974) Neural Basis in Mammals

• Cerebellum – Purkinje cells • Inhibitory connection to interpositus nucleus – Interpositus nucleus • CR output pathway • Error correction • Brain Stem – Pontine nuclei (CS) • Specialized sensory processing – Inferior Olive (US) • Activates interpositus nucleus and Purkinje cells Neural Bases of CC

A simpler diagram of Rabbit eyeblink conditioning Cerebellum

• Electrical activity

Purkinje cells

• Stimulating the inferior olive – Even specific tones, lights, etc. – Substitute for actual US • CC impaired after damage CS Modulation

• Latent inhibition not explained by RW

• Mackintosh (1975) – Salience of Sensory cues – Hippocampus • Animals without do NOT show latent inhibition Back to Aplysia CC in Aplysia

1. Aplysia and Eric Kandel Neural Bases of CC

In the long run, it’s all just LTP Proteins

• Two routes to long lasting memory – Activate CREB-1 • Synaptic growth – Deactivate CREB-2 • Rapid learning Addiction and Tolerance

• Homeostasis and compensatory responses – Environmental cues as CS

• Reducing reliance on drugs?

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