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Learning and Memory: Complete
Learning and Memory: Complete
L
et me begin by telling a little story. because we practice–repeat something over
When I was a graduate student we had and over. A further definition says,
to take an exam that Cornell does in “[Learning is] either a case of differential
an interesting way. They put you in a swivel- strengthening of one from a number of
chair surrounded by your committee responses evoked by a situation of need, or
composed of 4-5 faculty members. You are the formation of receptor-effector
spun around, and a question comes from the connections de novo; the first occurs
direction you face when you stop. One of the typically in simple selective learning and the
questions I was asked was, “What will be second in conditioned reflex learning" (Hull,
the most important accomplishment in the 1943). It is the strengthening of existing
field of neuroscience in the next 10 years?” responses and the formation of new
Knowing that knowledge usually advances responses to existing stimuli that make this
in small steps, I said that we would definition unique. So what is learning? It
gradually know more about most aspects of isn’t clear that we have an inclusive
the field, but I didn’t expect any major definition. It appears that learning is the
breakthroughs. To a man, the committee strengthening of existing responses or
agreed that in 10 years we would have a formation of new responses to existing
complete solution to the problem of learning stimuli that occurs because of practice or
and memory. The word complete was their repetition. How much practice? Sometimes a
term, presumably meaning that we would single practice session is sufficient as in
know everything there was to know about avoidance of painful or noxious stimuli.
learning and memory. Well, at the 10-year Sometimes a lot of practice is necessary as
mark, I sent them all a letter saying, “I told in learning to drive a car.
you so!” Their predicted event didn’t What then is memory? Again
happen. However, we have made some according to Kandel (2000), ". . . memory is
progress; that progress is what this chapter is the process by which that knowledge of the
about. world is encoded, stored, and later
retrieved." By this definition, memory is not
Definitions a thing; it’s a process. Interesting! In another
What is learning? According to Eric definition, "Memory is a phase of learning . .
Kandel (2000) “Learning is the process by . learning has three stages: 1. acquiring,
which we acquire knowledge about the wherein one masters a new activity . . . or
world.” While this definition is erudite, it memorizes verbal material . . . 2. retaining
doesn’t help us much in knowing what to the new acquisition for a period of time; and
study. Another definition (Kimble, 1961), 3. remembering, which enables one to
"Learning refers to a more or less permanent reproduce the learned act or memorized
change in behavior which occurs as a result material. In a narrower sense learning
of practice," is a little better. It tells us that merely means acquiring skill . . ." (Sargent
learning is more or less permanent; it won’t & Stafford, 1965). From these definitions,
always be there, but often will. It also tells we see that memory has to do with keeping
us that this is something that happens “knowledge” someplace and then retrieving
it when it is needed. What we don’t see here when given only the first few letters of the
is that the “knowledge” doesn’t have to word. At the same time, they deny ever
come into consciousness. I have two having learned the word previously. Implicit
cars–one with an automatic transmission, memory is often further parceled as
one with a stick shift. I don’t have to bring associative and non-associative. There are
into consciousness the process for shifting two well-known types of non-associative
gears when I get into the car that requires me learning: habituation and sensitization.
to do that–I just do it! Habituation is a decrease in response to a
benign stimulus when the stimulus is
Types of Memory presented repeatedly. A dog will be aroused
There are actually two basic kinds of when a strange tone is played. If the tone is
learning and memory. One is declarative or played over and over, the dog will
explicit; the other is non-declarative or eventually no longer be aroused by the tone.
implicit. Knowledge of facts–what we know We say that it has habituated. This kind of
about places, things and people–and the learning makes sense; it is not efficient for
meaning of these facts is explicit memory. an organism to go on responding to a
These things must be recalled into stimulus that has no meaning. The other
consciousness to be used. Patients who have form of non-associative learning,
bilateral medial temporal lobe lesions have sensitization, is an enhanced response to
an inability to learn and remember items of many different stimuli after experiencing an
factual knowledge. They can’t remember intense or noxious one. For example, an
people that they met the day before. They animal responds more vigorously to a tone
can’t remember what they did the day of lesser intensity once a painfully loud tone
before. Some people will further parcel has been played. Here we say that the animal
explicit memories as episodic (we remember is sensitized.
events) or semantic (we remember facts). As These two forms of learning also
Kandel (2000) points out, in either case the interact. Once a response has been
content of all explicit memories can be habituated, it can be restored by
expressed by declarative statements such as sensitization. In this case, we say that the
“I was here yesterday” (episodic) and “The animal is dishabituated. As an example: a
hippocampus has something to do with habituated startle response to a noise can be
memory” (semantic). restored by strongly pinching the skin.
Implicit memory involves In non-associative learning, it is not
information about how to perform necessary that the animal learns to associate
something; it’s recalled unconsciously. We the stimuli involved (thus the name). For
use implicit memory in trained, reflexive example, the dishabituated animal does not
motor or perceptual skills. I know how to learn to associate the noise with the pinch.
drive my car; I know how to get to work. As we shall see shortly, this is the hallmark
The same people with bilateral medial of associative learning. Not all forms of non-
temporal lobe lesions can learn simple associative learning are as simple as
reflexive skills–they habituate and are habituation and sensitization. For example,
sensitized, they can be classically and we learn language by imitation of people
operantly conditioned (see later). They can who already speak. This involves no
learn certain perceptual tasks. For example, association of stimuli and is clearly more
they can recall a word learned previously complicated than habituation.
In associative learning, we “learn” conditioned stimulus (CS). The response to
that two stimuli are associated with each it (again salivation) is called the conditioned
other or that a response is associated with a response (CR). The UR and the CR are
given event or has a given consequence. usually similar but often not identical in type
Perhaps important in clinical considerations, or strength.
a person can also learn that an outcome is Initially investigators thought that
not associated with a response. So a person classical conditioning involved simply
may learn that what happens to him is not learning that two stimuli were
related to what he does. Two sorts of contiguous–that they occurred close together
associative learning have been well studied: in time, one after the other. Now we think
classical conditioning and operant that what the animals learn is
conditioning. Classical conditioning is well contingencies–that existence of something
demonstrated by Pavlov’s famous depends upon existence of something else.
experiment in which he presented meat With this in mind, it is possible to see that
powder to a dog, causing it to salivate. He simply learning that two stimuli were
repeated the presentation, and each time the contiguous could often lead to behaviors that
dog salivated. If he repeatedly rang a bell were maladaptive, with animals associating
just before presenting the meat powder (they environmental events that had no real
were paired), the animal came to associate relationship. On the other hand, the
the bell with the presentation of the meat existence of superstitious behaviors, even in
powder, and it would begin to salivate when humans, suggests that this does occur.
the bell was rung. In fact, for a while it It is tempting to think of extinction
would salivate if the bell was rung but no as an example of forgetting, but alas it is not.
meat powder was presented (they were The difference is that something new is
unpaired). After a while, the bell stopped learned during the process of extinction–the
predicting the presentation of meat powder animal learns that the CS is no longer a
for the dog, and it ceased salivating when it signal that the US is about to occur, rather it
was rung. (This process is called extinction.) is a signal that the US will not occur.
It should be noted that for classical In operant conditioning (sometimes
conditioning to occur the ringing of the bell called trial-and-error learning), a person or
must precede the presentation of the meat animal learns that it gets a reward if it does
powder, often by a certain critical interval of something. So, a pigeon learns that it gets
time (of the order of 0.5 sec). One way to food if it pecks at a certain key, but not if it
look at classical conditioning is to think of pecks at another. A rat learns that it can
the bell as becoming a signal that the meat avoid getting an electric shock if it presses a
powder is about to be presented. bar at a certain time. Presumably what the
In Pavlov’s paradigm, the meat animal learns is that one of its many
powder normally elicits salivation without behaviors (pecking or bar pressing) is
experimenter intervention (it is innate or followed by food. It is constitutional in
perhaps previously strongly learned), and it animals to repeat behaviors that lead to
is called the unconditioned stimulus (US). positive reinforcement (something pleasant
The response is called the unconditioned or the absence of something unpleasant) and
response (UR). The bell comes to elicit avoid behaviors that lead to punishment or
salivation only after it is repeatedly paired negative reinforcement.
with meat powder; so it’s called the
Neuroscience of Learning and
Memory
Sensitization
In sensitization, a stimulus to one
pathway enhances reflex strength in another.
An example, again taken from experiments
in Aplysia, is shown in Fig. 18-5. Again,
stimulation of the siphon leads the animal to
withdraw the gill by activating sensory
neuron 1, which in turn activates a
Figure 18-4 - Simplified neural circuits involved in
the habituation process in Aplysia. There are about 24
sensory neurons in the siphon; these are glutaminer-
gic. T hey synapse o n 6 motor neuro ns that inne rvate
the gill and various interneurons as shown. The
control condition is shown on the left, the habituated
condition o n the right.
Habituation
What happens in the nervous system Figure 18-5 - Sensitization is produced by applying a
to produce habituation? Experiments noxious stimulus to the tail of the Aplysia’s tail,
performed in Aplysia californica, the sea activated sensory neuron 2. This, in turn activates a
facilitating interneuron that enhances transmission in
slug, were designed to address this problem. the pathway from the siphon to the motor neuron.
Their results are shown schematically in
Fig.18-4. If the siphon of the animal is
stimulated mechanically the animal motoneuron. If the tail of the animal is
withdraws the gill, presumably for stimulated just before the siphon is, then the
protection. That action is known to occur withdrawal of the gill is quicker and more
because the stimulus activates receptors in forceful. The mechanism of this appears to
the siphon, which activates, directly or involve serotoninergic, axo-axonic synapses.
indirectly through an interneuron, the As shown in the figure, activation of the
motoneuron that withdraws the gill. This is a sensory receptors in the tail activates,
simple reflex circuit. All of this is shown on through sensory neuron 2, a facilitating
the left side of the figure. With repeated interneuron that excites sensory neuron 1 in
the pathway leading the gill withdrawal. It opens Ca channels. The end result is that
does this either at the cell body or at the activation of this 5HT pathway by tail
terminals of the sensory neuron on the stimulation causes more transmitter
motoneuron or the interneuron. The substance to be released by siphon
consequence of the sensitization process is stimulation, the resulting larger EPSP leads
to increase the size of the EPSP in the to a larger response by the gill.
motoneuron without increasing the response
of sensory neuron 1. This will cause a With only short-term tail stimulation,
greater response in the motoneuron and the sensitization will fairly quickly disappear
therefore a greater withdrawal of the gill. when tail stimulation ceases. However, the
How all this occurs is illustrated in sensitization can be made relatively
Fig. 18-6, which shows an axo-axonic permanent by repeated tail stimulation. This
synapse as might occur between the long-term sensitization (and also long-term
facilitating interneuron and sensory neuron habituation) occurs because there are
1. Serotonin (5-hydroxytryptamine or 5HT) structural changes that occur in the
is released by the presynaptic axon onto the presynaptic terminals (sensory neuron 1, for
postsynaptic axon where it binds to example). With sensitization, there is an up
to 2-fold increase in the number of synaptic
terminals in both sensory and motoneurons.
Alternatively, with habituation, there is a
one-third reduction in the number of
References