Learning Flashcards
Pavlovian conditioning
The acquisition of a new behavioural (or physiological)
response to a previously neutral stimulus as a result of
experiencing a predictive relationship between it and a
biologically relevant stimulus.
Unconditioned Stimulus (US)
A stimulus that has natural relevance
Conditioned Stimulus (CS)
A stimulus that gains its relevance through learning
Explain the Pavlovian experiment
Pavlovian conditioning was first described by the physiologist Ivan Pavlov (hence
“Pavlovian”). He was conducting experiments on the reflexive control of digestion in dogs
when he came across a confound in his experiment. Instead of salivating when presented with
food, the dogs were starting to salivate as soon as the experimenter walked through the door.
Pavlov hypothesized that the dogs had come to associate the entrance of the experimenter with
the imminent arrival of food. He went on to conduct an experiment in which the signal that
predicted the food was the sound of a bell. Just as he had predicted, after a number of pairings
the dogs began to salivate as soon as they heard the bell, suggesting that they had successfully
associated the bell with food. Because the change in the dog’s behaviour was conditional on
the learning experience, Pavlov named the signal (in this case the bell) the conditioned
stimulus (“CS”), while the outcome (in this case, the food) is called the unconditioned
stimulus (“US”) because its effect on the dog’s behaviour was not conditional on the learning
experience. He named the response to the CS after training (in this case, salivation to the bell)
the conditioned response (“CR”) because the ability of the CS to elicit this response was
conditional on the learning experience.
Pavlovian conditioning can be appetitive or aversive. i.e. an association can be formed
between a CS and a pleasant US (e.g. food) or a CS and unpleasant US (e.g. electric shock). In
both cases, there is an increase in the vigour of the CR with the strengthening of the association
between the CS and US, because whatever the natural response would be to the US (be that
approach, fear, disgust, salivation etc.) is now the response to the CS – i.e. is that which
becomes the CR.
Necessary Conditions for learning
(1)Awareness of the CS-US relationship
(2)Biological Preparedness
(3)Neurobiological Dissociations
(4)Temporal Contiguity
(5)Food Aversions
(6)Blocking
Awareness of the CS-US relationship
In animals, we are mostly only able to assess the degree to which learning has occurred by
investigating a chance in outward behaviour. Human predictive learning, however, can be
assessed by two measures: 1) a change in the behavioural (or physiological) response to the
CS; 2) a change in the cognitive expectancy of the US following presentation of the CS
(usually measured by verbal self report). Often there is good concordance between these two
measures, raising the question of whether there is a causal relationship between cognitive
expectation and the acquisition of the behavioural response. Lovibond (Psychophsiology 1992
29 621) demonstrated this concordance. He paired some pictures of plants with electric shock
(CS+) and presented other plant pictures alone (CS-). CS+ and CS- trials were intermixed,
and participants were asked to rate the likelihood of shock to each CS. Those who rapidly
learnt this discrimination additionally showed a skin conductance response (SCR) on CS+
trials. No SCR was observed in participants who were unaware of the relationship between
CS and US.
Evidence that the cognitive expectation causes the SCR comes from the fact that when
Hugdahl and Ohman (J. Exp. Psychol: Human Learning and Memory, 1077 3 608) instructed
participants that there would be no more shock in an extinction phase (repeated presentation
of the CS without the US). In extinction, responding typically declines steadily across trials.
However, instructed extinction resulted in the immediate disappearance of the SCR. This is an
example of explicit learning, whereby awareness of the relationship between CS and US
causes the presence or absence of the behavioural response.
Biological Preparedness
There are some demonstrations of Pavlovian conditioning which appear to result from
implicit learning. For example, Hugdahl and Ohman also used fear relevant stimuli such as
snakes and spiders as CS+ and CS-. In contrast to the fear irrelevant stimuli used before
(flowers, houses), instructions to the participants that these stimuli would no longer be followed
by shock had little impact on extinction. This appears to be evidence of Implicit learning:
behavioural (or in this case physiological) change that occurs independent of cognitive
expectation. This type of learning may be facilitated in situations when the CS is highly
biologically relevant – for example signalling a potentially dangerous aversive US.
Neurobiological Dissociations
Bechara et al (Science 1995 269 115) studied fear learning in three patients with different
types of brain lesions, all involving bilateral damage to subcortical structures in temporal lobe:
the amygdala (AMG), the hippocampus (HC) and both AMG and HC damage (AMG+HC).
The AMG and AMG+HC patients failed to acquire an SCR response to a stimulus paired with
an aversive US (electric shock), whereas the HC patient showed normal conditioning.
However, when questioned afterwards about which CSs were or were not followed by the US,
the patient with AMG damage was able to report the associations accurately whereas the
patient with HC damage and HC+AMG damage were not. These findings demonstrate a
double dissociation between implicit SCR learning mediated by AMG, and explicit cognitive
learning mediated by HC.
Temporal Contiguity
One factor that determines learning is the temporal contiguity (closeness in time) of the
events involved in the learning episode – the longer the interval between the events, the less is
learned about their relationship.
Food Aversions
A common phenomenon that breaks the “rule” of temporal contiguity is conditioned food
aversion, whereby the taste of the food (or drink) and subsequent nausea are separated by an
interval of hours. Andrykowski and Otis (Appetite,1990, 14 145) interviewed patients about
types of food consumed prior to a chemotherapy session and took preference ratings. Many
patients developed aversions to new foods tasted prior to chemotherapy administration –
However, there was no relationship between conditioned food aversions and a) time between
that food consumption and chemo b) time between that food consumption and vomiting. Thus,
temporal contiguity is not always necessary.
Blocking
Contiguous pairings of events are also not always sufficient to bring about learning. This is
illustrated by the phenomenon of blocking. Block experiments tend to follow a particular
procedure: In stage 1, a stimulus (A) is paired with a US in stage 1 (this would be notated as
“A+”) while another stimulus (B) is not (“B-”), and this happens over repeated trials. In a
subsequent stage, A is paired with a new stimulus (X) and B is paired with another new
stimulus (Y). Both compounds are paired with the US (“AX+” and “BY+”) over multiple trials.
X and Y are then presented alone, and the behaviour of the participant observed. What is
normally found is that conditioned responding is seen to Y, but not to X. That is to say, the
participant has learned about Y, but not about X, despite having equal exposure to both. The
presence of A along with X in the second stage blocks an association forming between X and
the US. This difference can also be seen in neurological indices of learning. In an fMRI
experiment by Tobler et al (Journal of Neurophysiology, 2006 95 301), activation changes in
ventral striatum were seen more in BY+ trials than in AX+ trials. Furthermore, they found that
activity in an anterior region of orbitofrontal cortex during learning BY+ trials correlated with
the degree of behavioral difference.
Blocking can be thought of as resulting from predictive learning. In the first stage, over
multiple trials, A became a perfect predictor of the outcome. In the presence of A, X is not
learned about in the second stage, because the US is already fully predicted by A. This
“prediction error” account suggests that learning occurs when there is a discrepancy between
how much a US is expected (given the CSs present) and whether or not the US actually occurs.
We will explore this idea more in the next lecture.
Neural Basis of Pavlovian Conditioning
There is evidence that the dopamine (DA) system plays an important role in the acquisition of
associations, particularly when the learning is appetitive. In particular, midbrain areas such as
the ventral tegmental area (VTA)and Substancia Nigra (SN) have been the focus of much
research. Animals will work for intracranial self-stimulation (ICSS) of these areas (Olds &
Olds, 1963), and the administration of dopamine agonists increases rate of responding during
ICSS while dopamine antagonists attenuate responding (Gallistel & Karras, 1984).
Dopaminergic neurons in the substantia nigra and ventral tegmental area (VTA) in the brain
stem project to many forebrain structures, particularly the striatum, where they may facilitate
neural processing by releasing DA as a neurotransmitter. The VTA can be thought of as
projecting mostly to the ventral striatum, while the SN projects to the dorsal striatum.
O’Doherty et al (Neuron 2002 33 815) conditioned (trained) people to expect an appetitive
glucose solution following one picture CS and an aversive salt solution to another CS. They
conducted fMRI imaging, during the presentation of the two CSs.
The substantia nigra and VTA were more active during the CS associated with the appetitive glucose US relative to the CS paired with the aversive salt US.
Interestingly, the ventral striatum was more active during the anticipation of the reward than the receipt of it. There have been many different theories as to the exact role of dopamine in learning.
One hypothesis is that dopamine mediates the
hedonic or reward value of a stimulus (Wise, 1985).
Alternatively, the dopamine system may
be involved in incentive motivation, and could play a role during the anticipation of reward that corresponds to a motivational state of wanting or craving (Berridge, 1996).
Another hypothesis
is that dopamine functions as a prediction-error during reward learning (Schulz et al., 1997).
Evidence for the latter hypothesis comes from single cell neurophysiology recordings in which
(in the absence of a learned predictive cue), dopamine neurons have been found to respond to
the delivery of the reward itself, but after learning about a predictive cue, the neurons shift their
responses and respond instead to the presentation of the cue (Mirenowicz and Schultz, 1994)
Mechanisms of associative learning
Rescorla-Wagner rule.
Expectation and Surprise
Learning requires that the occurrence of the US is unexpected or surprising. Learning proceeds
in a negatively accelerated curve: with each CS-US pairing, expectation of the US increases
and surprise decreases. This is captured by an elegant and simple learning algorithm – the
Rescorla-Wagner rule. This rule states that increases in the associative strength of the CS (i.e.
the amount of expectation) results from the degree to which current associative strength
deviates from perfect learning (i.e. the amount of surprise). This deviation is known as prediction error. The Rescorla Wagner rule is captured by the following equation:
Excitatory and Inhibitory associative strength
Contiguous pairings of events are not sufficient to bring about learning (see the example
of blocking in the previous lecture). It also requires that the occurrence of the outcome is
surprising or unexpected. The Rescorla-Wagner rule formalizes this principle. It accounts
for acquisition by the CS of both excitatory and inhibitory associative strength. Thus, the
change in associative strength of a CS is determined by the discrepancy between perfect
prediction () and the sum of the current associative strength of all CSs present on that
trial (∑V).
When the US is present, =1, and the associative strength is excitatory (anticipation of the
presence of a US). When the US is absent, =0 and presents the opportunity for inhibitory
associative strength to accrue (anticipation of the absence of a US). According to
Rescorla-Wagner, this happens when a CS (e.g. CSB ) is presented with an excitatory CS+
(e.g. CSA) that has been fully learnt about previously, and no US is presented. Under these conditions, is zero (no US), but ∑V is positive because the presence of CSA predicted
that there would be a US. Therefore CSB will acquire inhibitory associative strength
(predicting the absence of a US that you’d otherwise expect to be there).
Lovibond et al (Behaviour Research and Therapy, 2000 38 967) demonstrated that a
conditioned inhibitor (E-) protected a conditioned excitor (C+) from extinction in human
participants. In the first phase, A was paired with shock on some trials (A+) and a
compound of AE was presented alone (without shock, AE-). This provides the conditions
for inhibitory associative strength to accrue to E. The control stimulus, B-, was never
paired with shock or an excitatory CS, and therefore neither predicted the presence or
absence of the US. Two more stimuli C+ and D+ were also trained as conditioned
excitors. Following this training, C and D were presented in an extinction phase (no US).
However, C was presented in a compound with E (CE-), while D was presented alone.
The following test phase showed that participants produced both SCR responding and an
expectation of the shock following C, but no response to D. Thus, the presence of the
conditioned inhibitor E with C in the extinction phase fully predicted the absence of
shock, so protecting C from losing excitatory associative strength.