Updated for 1999
Lecture Notes - Week 10
Learning - Chapter 14
TYPES OF LEARNING
1) Perceptual learning – ability to learn to recognize
stimuli that have been seen before
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Primary function is to identify and categorize objects and situations
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Changes within the sensory systems of the brain
2) Stimulus-response learning – ability to learn to perform
a particular behavior when a certain stimulus is present
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Establishment of connections between sensory systems and motor systems
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Classical conditioning – association between two stimuli
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Unconditioned Stimulus (US), Unconditioned Response (UR), Conditioned
Stimulus (CS), Conditioned Response (CR)
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Hebb rule – if a synapse repeatedly becomes active at about the
same time that the postsynaptic neuron fires, changes will take place in
the structure or chemistry of the synapse that will strengthen it (see
Figure 14.1)
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Rabbit experiment – tone paired with puff of air
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Instrumental conditioning – association between a response and a
stimulus; allows an organism to adjust its behavior according to the consequences
of that behavior
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Reinforcement – positive and negative
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Punishment
3) Motor learning – establishment of changes within the motor
system
4) Relational learning – involves connections between different
areas of the association cortex
5) Spatial learning – involves learning about the relations among
many stimuli
6) Episodic learning – remembering sequences of events that we
witness
7) Observational learning – learning by watching and imitation
other people
LONG-TERM POTENTIATION
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long-term increase in the excitability of a neuron to a particular synaptic
input caused by repeated high-frequency activity of that input
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hippocampal formation - specialized region of the limbic cortex
located in the temporal lobe. It contains:
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entorhinal cortex whose axons grow toward the dentate gyrus,
forming the perforant path
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dentate gyrus projects to pyramidal cells in CA3
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pyramidal cells project both to CA1 and to basal forebrain
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associative long-term potentiation - produced by association in
time between 2 sets of synapses; weaker strengthens after being paired
with stronger synapse
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series of pulses delivered at a high rate all in one burst will produce
long-term potentiation, but not the same number of pulses given at a slow
rate
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there are aftereffects which serve to prime future pulses by depolarizing
the postsynaptic membrane
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long-term potentiation requires two events:
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activation of synapses
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depolarization of the postsynaptic neuron
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NMDA receptors - type of glutamate receptor, critical in long-term
potentiation
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found in hippocampus, mostly CA1
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controls a calcium ion channel which normally is blocked by a magnesium
ion
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even the channel is stimulated by glutamate, calcium ions can’t get past
the magnesium
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but, if the membrane is depolarized, then the magnesium is ejected and
the channel can admit calcium ions
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therefore, need both glutamate and depolarization to admit calcium
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calcium is critical for long-term potentiation - both necessary
and sufficient
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entry of calcium activates some calcium dependent enzymes
1) protein kinase C (PKC) - normally in cytoplasm, activated
by calcium to increase synaptic transmission
2) CaM-KII - when activated by calcium it remains active even
after calcium is gone, until deactivated by another enzyme
3) tyrosine kinase - also plays a role in long-term potentiation
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nitric oxide - soluble gas used as a messenger in various parts
of the body
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produced by nitric oxide synthase in postsynaptic cell, communicates
with presynaptic terminal buttons - retrograde effect
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dendritic "spikes" - what are they? How do they happen? Why are
they significant?
AMPA receptors - control sodium channels - involved once long-term
potentiation has occurred
Long-term depression - low-frequency stimulation of the synaptic
inputs to a cell can decrease their strength; opposite of Hebb rule - weak
synapses not associated with strong ones become weaker
PERCEPTUAL LEARNING
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Involves learning about things, not what to do when they are present
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Simple perceptual learning, recognizing stimuli, takes place in appropriate
regions of sensory association cortex
1) Visual Learning
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inferior temporal cortex - necessary for visual pattern discrimination,
receives info from visual cortex
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ventral/dorsal streams - what and where
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delayed matching-to-sample task - requires that stimulus be remembered
for a period of time
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"remembering" the stimulus involves a neuronal circuit; it is the circuits,
not the individual neurons that recognize particular stimuli
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lesions of inferior temporal cortex disrupts an animal’s ability to remember
what it has just seen
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electrical stimulation of the inferior temporal cortex during the delay
causes forgetting
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responses of single neurons in inferior temporal cortex recorded when pairs
of stimuli shown
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found that when stimuli are paired, the neural circuits responsible for
recognizing them become linked together
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perception of either stimulus activates both circuits
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visual long-term memory involves the establishment of new circuits in the
inferior temporal cortex by means of synaptic changes
2) Auditory Learning
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auditory learning tasks modify response characteristics of neurons in various
parts of the auditory system
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study pairing tones with shock
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pretraining, neuron responds best to 9.5 Hz tone
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CS is 9-Hz tone, paired with shock
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after training, neuron now responds best to 9-Hz tone, and less to 9.5
Hz
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medial division of the medial geniculate nucleus (MGm) - important
for classically conditioned emotional responses
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receives info from auditory and somatosensory systems
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directly connected to the central nucleus of the amygdala
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which activates the nucleus basalis
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nucleus basalis contains acetylcholine neurons which innervate auditory
cortex, telling it to pay particular attention to the ventral division
of the medial geniculate nucleus (conveys auditory information)
S-R LEARNING
1) Classical Conditioning
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central nucleus of the amygdala involved in classically-conditioned emotional
responses
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see figure 14.29 -MGm eceives auditory and somatosensory info
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pairing of tone and footshock increases responses to CS
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extinction - somehow, NMDA receptors again involved - most likely
due to long-term depression
2) Instrumental Conditioning and Motor Learning
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2 pathways exist between sensory association cortex and motor association
cortex:
1) direct trascortical connections - short-term memory, and
with hippocampus involved in episodic memory
2) via basal ganglia and thalamus - used once no longer "new"
learning; Parkinson's example
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premotor cortex - monkeys raising arms for food; humans learning
motor task
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reinforcement:
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reinforcing brain stimulation discovered by Olds and Milner in 1954
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medial forebrain bundle - best place for self-stimulation; also
involved in natural reinforcers, such as food, water, or sex
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dopamine - likely serves as a neuromodulator, involved in reinforcement
- receptors in nucleus accumbens
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conditioned place preference - animals prefer to be where they have
encountered a reinforcing stimulus
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this process used to test influence of dopamine on reinforcement
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if given dopamine antagonist during place training, don’t develop conditioned
place preference
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also involved in negative reinforcement - dopamine antagonists prevent
avoidance learning
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amphetamine is a dopamine agonist - animals will work to get injections
of it
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drug discrimination procedure - train animals to press a certain
lever to receive food after it has been given a drug and to press another
lever after it has been given saline
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found that rats would press drug lever only if amphetamine put directly
into nucleus accumbens, rather than other brain structure
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they pushed the saline lever if they also had been given a dopamine receptor
blocker with the amphetamine into the nucleus accumbens
REINFORCEMENT SYSTEM
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must be able to first detect and then respond to reinforcement opportunities
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ventral tegmentum area - reinforcing stimuli activate neurons here,
which they stimulate release of dopamine in other systems
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neurons activated by both natural and conditioned reinforcers
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information received from 3 sites:
1) amygdala - involved in detection of CS for reinforcement
- if monkeys trained that food follows a visual stimulus, then amygdala
lesioned, the monkey forgets the association
2) lateral hypothalamus - neurons become active when monkeys
see food, but only when hungry - neurons show sensory-specific satiety;
activity related to presence of reinforcing stimuli
3) prefrontal cortex - secretes excitatory glutamate, which triggers
bursts of dopamine to be released from neurons in the ventral tegmental
area into the nucleus accumbens; may serve as monitor for reinforcement-seeking
activity
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a reinforcement system is required by instrumental conditioning
1) discriminative stimulus activates weak synapse
2) circumstance that causes animal to press lever activates a strong
synapse
3) if behavior is reinforced, then neurotransmitter/neuromodulator released
(dopamine), causing synaptic changes, strengthening weak synapses