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Thad A. Polk, PhD
Arthur F. Thurnau Professor of Psychology
University of Michigan
PROFESSOR BIOGRAPHY i
Professor Polk regularly teaches large lecture courses as well as small
seminars on topics ranging from the human mind and brain, to
cognitive psychology, to computational modeling of cognition. His
teaching at the University of Michigan has been recognized with
numerous awards, including the Excellence in Education Award
from the College of Literature, Science, and the Arts and the Arthur
F. Thurnau Professorship, the university’s highest undergraduate
teaching award. He was also featured in the University of Michigan’s
Professors Reaching Out for Students (PROFS) lecture series and was
named to The Princeton Review’s list of the Best 300 Professors in
the United States.
Professor Polk’s other Great Courses are The Aging Brain and The
Addictive Brain.
I’d also like to thank my lab manager, Holly Gagnon, who did a lot
of work tracking down articles and editing lectures as I worked on
the course. She played a particularly important role in the preparation
of the extensive bibliography. Thank you, Holly!
ACKNOWLEDGEMENTS iii
Table of Contents
Introduction
Professor Biography . . . . . . . . . . . . . . . . . . . . . . . . . . i
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . iii
Course Scope . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Guides
1 Learning 101 . . . . . . . . . . . . . . . . . . . . . . . . . . 4
5 Semantic Memory . . . . . . . . . . . . . . . . . . . . . . 41
Supplementary Material
Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
Image Credits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
TABLE OF CONTENTS v
vi The Learning Brain
The Learning Brain
One of our most important assets as human beings is our ability to
learn and adapt. As babies, we learn to crawl, stand, and eventually
walk. As toddlers, we learn to speak and understand our native
language. As we get older, we learn how to read and write; we learn
about history and literature, and about science and engineering.
We’re also constantly acquiring new skills, such as playing a sport or
mastering a musical instrument.
But how? This course will explore what’s going on in our minds
that allows us to learn so effortlessly and in so many ways. The
course will examine not only the psychological mechanisms, but also
the neural mechanisms, that are involved. You will discover that
your brain is equipped with multiple, powerful, but very different
learning mechanisms that depend on specific neural circuits and
that understanding how learning works can make you a more
effective learner.
COURSE SCOPE 1
somewhat unreliable, reconstruction of what might have happened.
Unfortunately, we usually can’t tell the difference between reality
and our reconstructions.
Next, the course will take you inside the skull to examine what’s
going on in your brain when you’re learning explicit information.
You will discover that one brain area, called the hippocampus, plays a
particularly important role, but that the information you learn tends
to get consolidated into other brain regions and therefore depends
less and less on the hippocampus as time goes by.
Next, the course turns to working memory and explores the critical
role it plays in all complex thought. You will learn about leading
theories of how working memory works, at both a psychological
and a neural level. You will also explore recent research examining
whether it’s possible to expand working memory and perhaps even
increase intelligence.
Throughout, the course will repeatedly identify specific ways that you
can apply what you’re learning to become a more effective learner in
your daily life. For example, you will discover study strategies that
are scientifically proven to improve learning and memory as well as
practice strategies that have been proven to help people learn skills
more effectively.
By the end, you will have a new understanding about how learning
works, how it is implemented in the brain, and how you can be a
more effective learner.
COURSE SCOPE 3
LECTURE 1
Learning 101
•• First, there are situations in which you learn something but there is
no opportunity to show it in your behavior. For example, suppose
that you learn some interesting but obscure fact about Berlin, such
as that a former airport there has been turned into a park. You
may have learned that fact and stored it in your memory, but it’s
quite possible that it will never influence your subsequent behavior,
unless you happen to visit Berlin or have a conversation about its
former airport.
•• But here is still at least one problem with our new definition. When
people think of knowledge, they typically think of information
that is consciously available and verbalizable, such as the fact that
birds have wings. We obviously do learn that kind of information,
but we also learn a lot of behavioral responses that are not
consciously available.
SUGGESTED READING
QUESTIONS TO CONSIDER
A Case of Amnesia
•• Henry Molaison was born in 1926 and grew up near Hartford,
Connecticut. When he was around 7 years old, he was run over
by a bicycle, hit his head, and was unconscious for a few minutes.
Later, he started experiencing epileptic seizures. At first, the
seizures were manageable, but they grew worse over time.
•• By the time Henry was 27, he was having about 10 seizures a day,
and large doses of anticonvulsant medication weren’t helping. He
was therefore desperate to find an alternative treatment. That’s
when a renowned neurosurgeon named William Scoville suggested
surgery, which can often be an effective way to treat epilepsy when
medications don’t work.
•• The hope was that the surgery would help control Henry’s seizures,
and it did. After the surgery, he only had about 5 minor seizures a
month. Unfortunately, he also suffered a very serious side effect:
He became profoundly amnesiac. As a result, that surgery has
never been performed again.
•• This task is difficult, and at first, Henry was bad at it. But after
practicing the task repeatedly for 3 days, he could trace the figure
quickly and accurately. Henry’s performance was changing as a
result of his experience—in other words, he was learning.
•• When Milner returned for the second and third day of training,
Henry didn’t remember having ever done the task before. He was
much better at it than he was the previous day, but his learning was
all unconscious. Apparently, learning a motor skill is fundamentally
different than learning a person’s name or other conscious facts.
•• And it’s not just motor skill learning that is preserved in amnesia.
For example, in 1980, Neal Cohen and Larry Squire investigated
the ability of amnesiacs to learn a perceptual skill in which they
had to learn to recognize unfamiliar visual stimuli. Specifically,
they asked them to read mirror-reversed text as fast as they could.
As in Milner’s study, the participants practiced the skill for 3
straight days. A group of nonamnesiac control subjects practiced
the same task.
Amnesiacs can learn perceptual and motor skills just like everyone
else. Although they may not consciously remember ever having
performed the tasks before, their performance continues to improve
with additional practice. Another way of putting this is in terms
of knowing how versus knowing that. Amnesiacs can learn how
to perform a skill, even when they don’t know that they’ve ever
practiced it before.
•• Pavlov hypothesized that the dogs had learned that they were likely
to be fed soon after he came in, so they were anticipating the food
even before it arrived. He began to investigate the phenomenon
systematically and discovered that if he consistently paired the
sound of a bell with the presentation of food, then pretty soon the
dogs would learn an unconscious association between the bell and
the food. In particular, they would salivate whenever they heard
the bell, even if there wasn’t any food.
SUGGESTED READING
QUESTIONS TO CONSIDER
1 The ants in the kitchen ate the sweet jelly which was on the table.
2 The warm breeze blowing from the sea stirred the heavy
evening air.
3 The rock which rolled down the mountain crushed the tiny hut
at the edge of the woods.
4 The old man resting on the couch read the story in the newspaper.
•• They also varied how many ideas from the original sentence these
simpler sentences conveyed. For example, the sentence “The jelly
was on the table” basically conveys a single idea, while the sentence
“The ants in the kitchen ate the jelly” conveys 2: that the ants were
in the kitchen and that they ate the jelly.
•• Subjects gave almost exactly the same ratings for new sentences
as they did for old sentences. Essentially, they had no idea which
sentences were new and which were old. This doesn’t mean that
they didn’t learn anything, but what they learned was the ideas
from the original sentences, not the exact wording.
•• Leo Standing has probably done the most work investigating how
much visual information we can remember, and his results suggest
that although we may forget a little of the visual information
we encode, there always seems to be more room to store new
information, particularly for pictures.
Previous Knowledge
•• How does what you already know affect your ability to learn new
information? It turns out that your previous knowledge can have
a dramatic effect.
•• If you don’t know what that paragraph was about, then it probably
sounded pretty weird and may have been difficult to make sense
of. But what if you know that the paragraph was about washing
clothes? Once you have that prior knowledge, everything in the
paragraph makes sense.
2 Subjects were told that the paragraph was about washing clothes
before they heard it. They understood the appropriate context
when they were hearing the paragraph.
3 Subjects were also told that the paragraph was about washing
clothes, but they were only told after they had already heard
the paragraph. They did, however, know what the paragraph
was about before they had to try to remember it.
•• After encoding, it’s too late. If you didn’t know what the paragraph
was about when you were encoding it, then you wouldn’t be able
to connect the new information to everything you already know
about laundry. You couldn’t build those associations with your
existing knowledge. And if those associations didn’t get built
during encoding, then they won’t be available when you try to
retrieve the information from memory.
•• When you get to the grocery store, you can walk through the
building in your mind’s eye and conjure up the strange image that
you stored there. You remember the cow being milked in your
living room, so you buy milk. You continue mentally walking
through each room in the building, retrieving the associated image
and recalling the next item on the list.
•• This age-old method has been proven to work time and time
again—because it exploits the principles in this lecture: You are
storing visual information rather than verbal information; storing
vivid, striking visual images rather than standard, ordinary images;
and connecting the information that you’re trying to learn with
information you already know.
QUESTIONS TO CONSIDER
2 What are some strategies you can employ in your daily life to
optimize your explicit learning?
T
HIS LECTURE IS ABOUT HOW SOME OF THE
INFORMATION THAT WE LEARN IS ACTUALLY
information that we infer. We are constantly
drawing conclusions about the information
that we encounter, and some of the
information that we store during learning
may be our inferences rather than accurate
memories. And this has important real-world
implications for eyewitness testimony and for
how to interview witnesses.
Making Inferences
•• We are constantly making inferences to construct a plausible
mental model of the situations we encounter. And we tend to
remember our mental simulation of events, including all the
inferences we made, rather than the original information itself.
•• And it turns out that we make these kinds of inferences all the
time. We do it when we’re encoding new information and when
we’re retrieving information from our memory. In an experiment
that demonstrates this fact very clearly, a group of subjects were
asked to try to remember the following story:
•• Then, all subjects were asked the same question: Do you remember
reading the sentence “She’s deaf, blind, and cannot speak?” That
sentence wasn’t actually in there, but the scientists found that
about half the group who were told that the story had been about
Helen Keller said that they remembered that sentence. In contrast,
only 5% of the control group said that.
•• But before that happens, they may or may not receive some
misinformation—some subtle information that could potentially
distort their memory for the original event. Then, everybody’s
memory gets tested and the researcher determines whether the
people who were misinformed exhibit memory distortions relative
to the people who weren’t misinformed.
•• Using the phrase “smashed into each other” suggests that it may
have been a more violent collision, while the phrase “hit each other”
doesn’t have the same strong connotation of a violent collision. But
that’s the only difference between the 2 groups.
•• The control group is asked this question: Did the car pass the
Datsun when it was stopped at the yield sign? The misled group
is asked a slightly different question: Did the car pass the Datsun
when it was stopped at the stop sign? After a delay, both groups
are shown a picture of the Datsun parked at a yield and another
picture of the Datsun parked at a stop sign. Then, they’re asked a
final question: Which one did you see?
•• In contrast, the people who read the neutral description that was
not misleading correctly picked the hammer 72% of the time.
So, there was a whopping 35% difference in accuracy between
the 2 groups. This is the standard misinformation effect seen in
previous studies.
•• Subjects chose the hammer 72% of the time, which was very
similar to the people who read the neutral description. These
results suggest that “hammer” is still in their memory somewhere.
They didn’t overwrite it; instead, they augmented their memory.
2 The goal of letting the eyewitness tell his or her story without
interruption is to try to eliminate the possibility of introducing
a misinformation effect. A police officer asking an eyewitness
about a traffic accident he or she just saw might unconsciously
say something that could bias the witness testimony. By
avoiding asking any questions at first, there is no way the
officer’s questions can bias the eyewitness’s memory.
SUGGESTED READING
QUESTIONS TO CONSIDER
•• These types of memories are not tied to a specific time and place
and are not remembered from a personal perspective. They are
just disembodied facts that you happen to know; they are part of
your general knowledge about the world.
•• The intuitive idea that many people initially have is that there are
some specific features that define a mental category. And if an
object has those features, then it’s guaranteed to be a member of the
category. If it doesn’t, then you know it’s not. So, these features are
both necessary and sufficient to determine category membership.
•• According to this view, you don’t just check off the defining
features and decide whether an object is guaranteed to be in or
out of the category. Rather, you ask how similar is it to other
members of the category.
SUGGESTED READING
QUESTIONS TO CONSIDER
•• The 2 temporal lobes are on the bottom of the left and right side of
your brain. They are below, or inferior to, the parietal lobes and are
in front of, or anterior to, the occipital lobes. The medial temporal
lobes—the part of each temporal lobe toward the middle of the
brain, away from the ears—are the parts of the brain that were
surgically removed in Henry Molaison (the amnesiac patient you
learned about in lecture 2), and doing so led to a profound, but also
quite isolated, deficit in learning explicit, conscious information.
•• It turns out that your memory of picking up and biting into that
apple depends on those same brain areas. When you remember
what the apple looked like, you’re activating the occipital cortex.
When you remember what biting it sounded like, you’re activating
the auditory cortex. When you remember what it tasted like, you’re
activating the insula—and so on.
•• Why is our memory system organized this way? Why not just
consolidate the memory into the cortex right away? Why does
consolidation take so long? Isn’t that inefficient?
•• One synapse might get a little bit stronger and excite the next
cell a little more than it did before. Another synapse might get
weaker and excite the next cell a little less. By changing those
synaptic strengths in just the right way, the brain learns a long-
term memory. And when appropriate cues are encountered in the
future, they will trigger the neural activation pattern associated
with that memory. That’s memory retrieval.
SUGGESTED READING
•• About half the questions on the test were basic factual questions
about the material. On these questions, all 3 groups performed
about the same. Once again, marking some of the text as
particularly important—in this case by underlining—didn’t
help. The other half of the questions required students to draw
inferences based on what they had read. On these questions, the
students who had used underlining during both the initial study
period and the review period actually performed worse than the
other 2 groups.
If you have read a chapter and then reread it, you will recognize
a lot of what you read. And that familiarity may fool you
into thinking that you really know what’s in the chapter.
Unfortunately, you may be in for a rude awakening if you’re then
tested on the chapter and have to actually recall the information.
This happens all the time in classrooms around the world. Students
who do poorly on an exam are often very surprised, because they
really thought they had mastered the material. And part of the reason
is probably because rereading gave them the illusion of mastery.
•• A final reason that rereading probably isn’t the best study strategy
is very simple. Other techniques have been demonstrated to be
significantly better. And any time you spend rereading is time that
you’re not spending studying in these other, more effective ways.
•• There are 2 other study techniques that were rated as even more
effective by Dunlosky’s review than generating explanations and
interleaving different topics when studying. The first is distributed
practice, which refers to spreading out your study over time with
breaks or lags between study sessions.
•• Then, Bahrick gave everyone a test 30 days after their last practice
session to see how much they remembered. The group that crammed
all their study into a single day remembered 68% of the Spanish
word translations. The group that had study sessions separated by
a day remembered 86% of the translations. And the group that
studied once a month remembered 95% of the translations.
•• Keep in mind that being tested on a passage meant that the students
didn’t get to restudy it, so they had actually studied those passages
for less time than the other passages, yet they learned more. This is
a very valuable lesson for students: Rather than rereading a chapter
they’ve already read, students would be much better off asking a
friend to test them on the material.
SUGGESTED READING
QUESTIONS TO CONSIDER
1 Do you have any personal study techniques that work well for you
but that weren’t discussed in this lecture? If so, why do you think
they’re effective, based on what you’ve learned in the course so far?
2 Are there any specific ways that you can apply what you learned in
this lecture to make your learning more effective?
•• This certainly seems like a reasonable idea. After all, people are
quite different in every way you can think of, so it makes sense to
think that people probably also learn in different ways. And if so,
then tailoring instruction to match someone’s learning style could
have real benefits.
•• And consistent with the intuition, the idea of learning styles has had
an enormous impact in education and in society more generally.
Testing to determine learning styles has been recommended by
several education organizations, and a number of textbooks in
educational psychology discuss the merits of the approach. The
idea of learning styles has also led to commercial products and
training programs that generate significant revenue.
•• But there is not yet solid scientific evidence to support the claim
that tailoring instruction to match learning styles works. It’s
possible that future research will demonstrate the value of assessing
learning styles and designing individualized instruction, but the
evidence to date has a number of problems.
2 A lot of the work in the field is based on what students report that
they like rather than what actually improves their learning. Most
of the tests that have been designed to assess learning styles ask
students to indicate their preferred way to perform different tasks,
and based on those preferences, the student gets categorized into
a particular learning style. But the critical issue isn’t how much
students like a particular approach, but rather how much they
learn using that approach. Learning methods that are actually the
most effective might not be what students most prefer.
•• Many people think that the answer to these questions is yes. Many
popular books tout the benefits of listening to classical music—
not only to improve cognitive function, but also to reduce stress
and heal the body. Entire companies have sprung up marketing
and selling products based on the assumed benefits of listening
to classical music.
•• This is the finding that has since been dubbed “the Mozart effect,”
although the scientists who conducted the study never used that
term in the original paper.
•• The popular press latched onto these findings because they seem
to suggest that the simple act of listening to 10 minutes of Mozart
can increase your IQ by almost 10 points. This is certainly an
interesting finding, but there are 2 important limitations that often
get glossed over:
•• On the other hand, there are also reasons to doubt the authenticity
of at least some recovered memories. First, a number of studies
have shown that people tend to remember emotionally arousing
information better than emotionally neutral information. For
example, many combat veterans want to forget what they witnessed
but are tormented by extremely vivid memories of the horrors they
experienced. If forgetting is a psychological defense mechanism,
then you might expect the opposite.
SUGGESTED READING
•• The opposite can also happen; that is, rather than learning to ignore
a stimulus, you can learn to become more sensitive to it. This is
called sensitization, and it’s also a form of nonassociative learning.
•• Pavlov devoted much of the rest of his career to studying this kind
of learning, which has come to be known as classical conditioning.
It’s also sometimes called Pavlovian conditioning in honor of
its discoverer.
•• Learned food aversions are quite common, and they can be induced
in laboratory animals very easily. For example, if rats are given a
novel food to eat and are then injected with a drug that makes
them sick, they will quickly learn an aversion to the novel food.
And they learn this aversion much faster than Pavlov’s dogs learned
their association. In fact, animals can learn food aversion after a
single trial. And they can learn it even if they don’t get sick until
hours after eating the novel food.
•• The critical test was what would happen if Thorndike kept putting
the cats back in the same box. And he found that they gradually
got faster and faster at escaping. After 5 or 6 trials, the cats learned
to go straight to the lever and push it, allowing them to escape
within just a few seconds.
QUESTIONS TO CONSIDER
•• Imagine that you are just starting to learn to play the piano and
that your first lesson is how to play a D major scale. The first time
you do this, you will be pretty slow. But the second time you do
it, you will probably be a little bit faster. And after about a week of
practice, you will be much faster. You’re not going to be a skilled
pianist after just a week of practice, but you will be dramatically
better than you were on the first day.
a Suppose that you’ve never played golf and that you take
a lesson. The golf pro will probably start out by giving
you some verbal instructions, such as “keep your left arm
straight when you swing.” Now you have a bunch of verbal
instructions that you will dutifully try to execute. In fact, you
may try to memorize some of the verbal instructions or even
write them down. And you may consciously and deliberately
rehearse the verbal, declarative knowledge that the pro told
you as you’re trying to hit a golf ball.
•• For example, when you walk, you’re not usually thinking about
how to walk; instead, it’s an automatic set of associations. That’s
what production rules are: automatic associations. Walking is not
conscious; it’s an implicit, unconscious, procedural skill.
•• This process continues for each of the other instructions for tying
shoelaces. You create separate production rules corresponding to
each piece of declarative knowledge.
SUGGESTED READING
QUESTIONS TO CONSIDER
•• First, they have to realize that not all sounds are language. They
need to distinguish language sounds—that is, speech—from all
the other sounds in the environment, such as the sounds of the
doorbell ringing and a dog barking. This is the first obstacle in
language acquisition: just figuring out what the target is. What
are the sounds that babies need to focus on to start learning a
specific language?
•• Let’s say that a baby has figured out that some of the weird sounds
that come out of his or her parents’ mouths correspond to a kind of
communication. And the baby begins focusing on them and trying
to figure them out. That’s a very far cry from actually understanding
what is being said, much less being able to produce language. And
before a baby can understand what it means, he or she needs to be
able to recognize the sounds that make up the words.
•• For example, when you hear the word “cat,” you need to somehow
chop this continuous stream of noises into component phonemes:
“cuh,” “ah,” and “tuh.” And even if you have done that, you also
need to figure out which phonemes are grouped together in the
same word and which phonemes are in different words. That’s
actually very difficult, because when people speak, all of the words
glide together. It’s just a continuous stream of phonemes, and it’s
not at all clear where the word boundaries are.
•• Worse yet, the same sound can mean different things, and different
sounds can mean the same thing. For example, the word “bank”
can be used to refer to a financial institution or to the side of a
river. Conversely, the words “car” and “automobile” refer to the
same thing. Such irregularities make the task of learning word
meanings very difficult.
•• But let’s say that you have distinguished language sounds from
other sounds, parsed language sounds into individual phonemes,
figured out where the word boundaries are, and even begun to
figure out what some of the words mean. You still have to figure
out the grammar of your language. How do you put different
words together into phrases? How do you combine phrases into
sentences in a grammatical way?
•• Children are learning a lot about language in the first few years
of life. And they do all of this without a lot of feedback. In other
words, adults don’t usually correct their children’s grammar or
pronunciation. Psycholinguists who have tried to correct their own
children’s grammar found that providing explicit feedback when
the child makes a mistake doesn’t help him or her acquire language
•• For example, all natural languages are based on words. And those
words are based on combinations of specific sets of phonemes.
Those words are divided into specific parts of speech, such as
nouns and verbs. And those parts of speech help determine how
words are put together into phrases and sentences.
Language Development
•• What do we see when we actually observe children trying to
acquire language? Let’s start at the level of just distinguishing the
phonemes of a language.
•• Research has shown that babies can hear the phonetic distinctions
in any natural language. But as they get more and more exposure to
their native language, their speech perception gets specialized to that
language. And sounds that correspond to different phonemes in that
language begin to sound more distinctive, while different sounds
that correspond to the same phoneme begin to sound more similar.
SUGGESTED READING
QUESTIONS TO CONSIDER
•• So, why did the snails behave any differently? Kandel and his
colleagues discovered that the neurons in the circuit reduced
the strength of their signals. Specifically, they released less
neurotransmitter than they did before habituation.
•• When those receptors get turned on, they might trigger the next
neuron to fire and send a neurotransmitter signal to another
neuron—and so on. That’s the way information gets sent through
the neural circuits in your brain. All of your perceptions, decisions,
and actions are based on neurons firing and triggering other
neurons to fire.
•• But Kandel and his colleagues found that after repeated gentle
touching, the sensory neurons in the siphon started releasing
less and less neurotransmitter. As a result, the receptors on the
motor neurons weren’t turned on as much, and the gill withdrawal
response became smaller and smaller. That was the neural basis
of habituation: reduced neurotransmitter release. And similar
findings have been reported in many other species.
•• When the monkeys first started doing the task, Schultz noticed
that the VTA neurons tended to fire when the monkey got a juice
reward. He saw a big spike of neural activity as the VTA neurons
fired and released dopamine. But once the monkeys figured out
that they would get the juice as long as they pressed the lever
when the light came on, then the VTA neurons stopped firing in
response to the juice.
•• More than 20 years after Hebb originally proposed this idea, Tim
Bliss and Terje Lømo discovered empirical evidence that Hebb was
right. Specifically, they found that if they artificially stimulated
a neural circuit with high-frequency electrical activity, the
strength of the synaptic connections in that pathway got stronger.
Furthermore, they found that the strength, or potentiation, of
those synaptic connections lasted for days or even weeks, so they
referred to the phenomenon as long-term potentiation (LTP).
•• A few years later, Gary Lynch and his colleagues demonstrated that
it was also possible to induce the long-term depression (LTD) of
synaptic connections. In that case, the synaptic connections get
weaker rather than stronger.
•• Normally, rats swim around until they stumble upon the platform
but then quickly learn its location and swim straight for it when
they’re put back in the pool. But after being injected with the
LTP-blocking drug, the rats were significantly impaired at learning
the platform’s location, suggesting that LTP is playing a role in
real-world learning.
•• LTP and LTD have been found in numerous parts of the brain,
but they have been most studied in the hippocampus—which is
crucial for explicit learning. These are basic neural mechanisms
that could play a role in both implicit and explicit learning.
SUGGESTED READING
Space
•• To maximize long-term learning, practice should be spaced out,
or distributed, over time rather than crammed into a short period.
•• One month after the final training session, all of the residents were
asked to perform the surgery on a live rat while experts who did
not know about the experiment observed. The residents whose
training was spaced performed much better. They all completed
the surgery successfully, while some of the residents in the other
group did not. Furthermore, their surgically repaired blood vessels
were more stable, and the experts rated their performance as being
significantly better.
Challenge
•• Research suggests that your practice will be more effective if
you deliberately challenge yourself. If you can identify specific
challenging areas to work on, set specific goals within those
areas that are difficult—but not impossible—to achieve, and
•• One potential explanation for why is that when you actually have
to perform the real skill later, you’ll typically have to perform all
the subparts in an unpredictable order. Using random practice
therefore better prepares you for the real execution of the skill. For
•• Spacing and randomizing can also be taken too far. For example,
spacing becomes ineffective if the delay between practice sessions
is so long that the learners have forgotten what they previously
learned. And although randomizing practice conditions is helpful,
there is no need to include conditions that you would never actually
encounter in real life. For example, although it’s helpful for a
basketball player to practice shooting from a variety of different
distances to the hoop, there is no need for the player to practice
shots from 200 feet away.
QUESTIONS TO CONSIDER
1 What new skill would you like to acquire? How might you apply
what you learned in this lecture to help you acquire that new
skill faster?
131
•• Although most addictions involve taking drugs, many scientists
believe that some other behaviors should also be considered
addictions. An example is pathological gambling, which involves
compulsive behavior and can have significant negative consequences.
•• The same is true for other drugs. Opiate drugs, such as heroin
and prescription painkillers, tend to produce a sense of euphoria
and also cause constipation. So, when they’re withheld, the addict
might experience withdrawal symptoms such as agitation and
diarrhea—the mirror opposite of the effects produced by the drug.
•• However, if you get the reward when you weren’t expecting it,
then there is a reward prediction error—you failed to predict the
reward when it showed up. And that means you should try to learn
so that you can better predict rewards in the future. That’s when
you learn new associations between whatever stimuli are in the
environment and the reward you just received.
•• How does all of this relate to addiction? All addictive drugs lead
to an artificially large burst of dopamine. And that large burst of
dopamine is what makes the drugs so addictive.
•• Notice that the reward for drug addicts actually isn’t unexpected.
If they are chronic users, they know very well what to expect from
taking their drug of choice. In fact, they’ve probably developed a
tolerance to their drug so that the rewarding effect isn’t as strong
as it used to be.
•• But the brain doesn’t know that—so it behaves the way it always
does when it gets a dopamine burst. It interprets it as meaning
that an unexpected reward is on its way and does 2 things: First,
it triggers wanting or motivation. (This is not regular wanting, but
the pathological craving that addicts report feeling.)
QUESTIONS TO CONSIDER
•• You might think that evidence like that proves that long-term
memory and working memory depend on different underlying
systems. But there’s a potential alternative explanation. Suppose
that the long-term memory tasks are just more difficult than the
working memory tasks. Then, you might perform worse on the
long-term memory tasks even if they depend on the same neural
systems as the working memory tasks.
•• You could imagine different ways of storing those words. You could
store images of what the words look like based on the shapes of the
component letters. You could also imagine storing representations
of the words’ meanings. But a key assumption of the phonological
loop is that you store the words in a sound-based format.
•• But what’s really interesting is that we only have trouble with items
that sound alike, not with items that look alike or items that have
similar meanings.
•• For example, suppose that you are asked to remember a few words
in your working memory, but instead of saying them out loud,
you are only allowed to read them on a piece of paper or on a
computer screen. The words are presented in a visual format, not
a phonological format.
•• The model assumes that you will use your articulatory loop to say
the words to yourself and convert them into a phonological format.
And that’s the format that they’re stored in. So, even though the
words were presented visually, they are stored phonologically.
SUGGESTED READING
QUESTIONS TO CONSIDER
•• For example, look around wherever you are right now and store
a visual image of what your environment looks like. Now close
your eyes and bring that visuospatial image to mind. That’s
your visuospatial sketchpad at work, temporarily storing that
visuospatial information while your eyes are closed.
•• Brooks found that the 2 response types had opposite effects on the
2 tasks. First, in the visuospatial working memory task, he found
that people are faster if they get to say “yes” or “no” out loud and
•• You might think that talking is just easier than pointing and that
people would always be faster when they get to say their responses.
But Brooks found the opposite pattern for the sentence task: People
were faster when they pointed to the response on a piece of paper
and slower when they spoke their responses out loud.
•• But there are also critical differences between the episodic buffer
and long-term episodic memory. First, the episodic buffer is short-
term, so information in the buffer must be rehearsed or it will
quickly fade away. In addition, the capacity of the episodic buffer
is assumed to be limited, whereas long-term episodic memory is
assumed to have almost unlimited capacity. In particular, the
episodic buffer is assumed to have a capacity of about 4 chunks
(or meaningful units) of information.
SUGGESTED READING
•• The holes were then covered, and a blind was lowered between the
animal and the holes. Then, there was a delay of anywhere from 15
seconds to a little more than a minute. After the delay, the blind
was removed, which allowed the animal to reach out through its
cage into one of the covered holes.
•• If it reached into the hole with the apple, then it got to eat the apple.
If it reached into the other hole, then the experimenters showed
the monkey where the apple really was but didn’t let the animal
have it. Monkeys are smart, and they quickly figured out the task
and began consistently reaching into the hole containing the apple.
•• While the monkeys were doing this, Fuster and Alexander were
recording from neurons in their brain. And they found a bunch of
neurons that fired continuously during the delay period and then
stopped firing after the animal made its response.
•• These neurons were in the prefrontal cortex. Recall that the cortex
is the thin layer of gray matter around the outside of the brain,
and the prefrontal cortex is the cortex in the front of the brain.
•• The part of the brain that is associated with the central executive
is the prefrontal cortex. According to Baddeley’s theory, we should
expect to see neural activity in the prefrontal cortex during working
memory tasks. And we do.
•• For example, one neuron was active whenever the monkey was
trying to remember that the square had been in the upper-right
corner of the display, and a different neuron was active when the
monkey was trying to remember that the square had been in the
lower-left corner. But the neural responses were consistent: The
same neurons always fired when the monkey was remembering
“upper-right,” and a different set was always active when it was
remembering “lower-left.”
•• And it turns out that it’s not just the prefrontal cortex that’s
involved in working memory. Subsequent experiments have
demonstrated neural activity during the delay period in a number
of more posterior parts of the brain. And the posterior brain area
that is activated is typically the area that represents the kind of
information being remembered.
•• One hypothesis for what these prefrontal regions are doing is the
idea that the prefrontal cortex provides top-down excitation of
posterior brain regions during working memory tasks.
The Convergence of
Psychology and Neuroscience
In Alan Baddeley’s model of working memory, which
assumes that working memory can be subdivided into
different components, he argues that we should distinguish a
phonological loop from a visuospatial buffer and that both of
these are controlled by a central executive.
That model actually maps pretty naturally onto the neural
results addressed in this lecture. For example, Jonides and Smith
found that storing verbal information depends on completely
different brain circuits than storing spatial information. And that
makes perfect sense, assuming that we use the phonological
loop to store verbal information while we use the visuospatial
sketchpad to store spatial information.
Furthermore, a natural interpretation of the neural data is that
the prefrontal cortex provides top-down excitation of posterior
brain areas. That seems similar to the top-down control that
you would expect Baddeley’s central executive to exert. And
Baddeley assumes that the central executive component of
working memory is implemented in the prefrontal cortex.
QUESTIONS TO CONSIDER
•• The other half of the children served as a control group. They also
performed the same working memory tasks for the same amount of
time, but the tasks were all easy and didn’t become more difficult
as the children improved. So, the only real difference between the
2 groups was in the intensity of the working memory training and
whether it stretched the children or not.
•• They also found another neural change that was specific to working
memory training: The connections between prefrontal regions and
posterior brain regions became stronger. How could strengthening
these connections help improve working memory performance?
•• But for that to work, you need good connections between the
prefrontal cortex and the posterior brain regions that it excites.
And sure enough, as those connections get stronger, working
memory performance improves.
SUGGESTED READING
QUESTIONS TO CONSIDER
•• For example, imagine a high school student who believes that she
is bad at math. Maybe she had a bad experience in an earlier math
class, or maybe other people have told her that she’s not the type
of student who does well at math. But for whatever reason, she
has now come to believe that she is just not cut out to learn math.
As a result of that low self-efficacy, she decides not to try. Rather
than risking failure, she chooses to take other, less challenging
courses instead.
•• But now imagine another student named Holly who also studied
hard for the same exam and also performed poorly. But unlike
Peter, Holly assumes that if she had prepared in a different
way, then she could have done better. So, maybe she goes to the
instructor’s office hours to review her exam, figure out what she
missed, and identify more effective ways to study in the future.
•• People with a fixed mindset view their talents and abilities as fixed
and unchangeable. The idea of a fixed mindset and the idea of low
perceived control are similar. If your poor performance is a result
of fixed abilities, then you have very little perceived control. And
from that point of view, putting a lot of energy into studying or
working hard doesn’t make a lot of sense. In fact, students with
a fixed mindset might view the need to work hard as a sign of
inferior intelligence.
•• In one of the sessions, half the students were paid a dollar for each
configuration they were able to reproduce. But in the next session,
they went back to doing it without any monetary incentive. The
other half of the students never got any money and just did it for fun.
Perceived Control
•• The fourth factor that has been shown to have a significant
impact on learning is the value that students place on what they’re
learning. Do students care about what they’re learning? Do they
think that what they’re learning is important in some way? If so,
that can significantly increase the amount that they learn.
Bandura, Self-Efficacy.
Deci and Flaste, Why We Do What We Do.
Duckworth, Grit.
Dweck, Mindset.
QUESTIONS TO CONSIDER
•• But the researchers found the opposite for words that were highly
arousing: 20 minutes after studying the pairs, the participants
remembered about 20% of the numbers associated with words
that were emotionally arousing. That’s about the same as for the
neutral words. But the next day, their memory for those numbers
actually got better rather than worse. Now they remembered
almost 40% of the numbers. And after a week, their performance
was even better.
•• More recent studies have found similar results. As time passes, the
gap between emotional memories and neutral memories grows.
And that has led many scientists to hypothesize that emotional
arousal strengthens memory consolidation.
•• Stress tends to make you narrow your focus of attention to the most
central, important information. And memory for that information
is often improved. But memory for other information is typically
impaired, precisely because you ignored it.
•• The rat wouldn’t want to walk down the same arm twice because
the peanut won’t be there anymore, so that would be considered
a mistake, while walking down an arm that the rat hasn’t visited
before would be considered a correct answer.
•• Cahill and his colleagues tested this idea by presenting people with
2 stories to remember. One was an emotional story about a boy
walking with his mom, getting hit by a car, and being rushed to
the hospital to be treated for life-threatening injuries. The other
was a neutral story in which the boy and his mom walk to the
hospital and observe a simulated emergency response.
•• For example, rats and other animals that have lesions to the
amygdala don’t exhibit normal fear conditioning. After such
lesions, these animals do not learn the standard association between
a conditioned stimulus, such as a tone, and a fearful conditioned
stimulus, such as a shock.
SUGGESTED READING
QUESTIONS TO CONSIDER
2 Can you think of any ways to reduce the level of chronic stress
in your life (and thereby potentially improve your learning and
memory)?
•• Aserinsky also noticed that people move their eyes behind their
closed eyelids during these periods of significant brain activity,
but not during other periods of sleep. He therefore referred to the
highly active periods as rapid eye movement (REM) sleep and to
the other periods of sleep as non-REM sleep.
•• During slow-wave sleep, our core temperature drops, and it’s more
difficult to wake up. We typically spend 20 to 40 minutes in this
kind of deep sleep before transitioning back into lighter non-REM
sleep for a few minutes and then finally into REM sleep.
•• The first period of REM sleep is usually pretty brief, lasting only
1 to 5 minutes. Then, we transition back into non-REM sleep,
and the cycle begins again. Overall, we spend about a quarter of
our total sleep time in REM sleep, about half in light non-REM
sleep, and another quarter in deep, slow-wave non-REM sleep, but
the proportion changes throughout the night.
•• With each new cycle, the REM periods tend to get a little bit
longer, while the deep, slow-wave periods tend to get shorter and
may disappear entirely in the last few cycles. As a result, the first
few cycles are dominated by deep, slow-wave sleep, while later
cycles are dominated by REM and light non-REM sleep.
•• The fact that the first few hours of sleep are qualitatively different
from the last few hours has led some scientists to hypothesize
that different types of sleep help to consolidate different types
of learning. Specifically, they have hypothesized that slow-wave
sleep is more important in the consolidation of explicit, declarative
memories, while REM sleep is more important in the consolidation
of implicit, procedural memories.
•• Björn Rasch, Jan Born, and a few colleagues did just that in one
of the most creative sleep experiments, in which human volunteers
learned an object location task before going to sleep for the
evening. Specifically, they were presented with an array of cards
•• But while they were learning the locations of the cards, they were
also presented with a smell—specifically, the smell of roses. Smells
can be very strong memory cues, and in this case, the scientists
used the smell of roses to try to reactivate memory of the card
location task during sleep. So, they monitored the participants’
brain waves while they were sleeping and then presented the smell
of roses during different periods of sleep.
SUGGESTED READING
Hobson, Dreaming.
Stickgold and Walker, The Neuroscience of Sleep.
Walker, Why We Sleep.
QUESTIONS TO CONSIDER
2 Are there any specific steps you can take to improve the quality of
your sleep?
•• Around the year 2000, Denise Park and her team gave people tests
that tap into fluid processing ability—such as asking people to try
to remember random words or line drawings or testing how fast
they could perform tasks that didn’t depend on knowledge—and
generated plots of people’s performance as a function of their age.
•• Park’s group also tested this kind of processing ability, using tasks
that tested things like world knowledge and vocabulary. The
results on these tasks looked very different than the results for
fluid intelligence. Here, the older groups tended to perform better
than the younger groups.
•• Recall that the prefrontal cortex is kind of like the CEO of the
brain, controlling what we pay attention to, managing goals, and
delegating responsibilities to other brain processes. In particular,
the prefrontal cortex is hypothesized to play a central role in
working memory. When Raz and his colleagues plotted prefrontal
cortex volume versus age, they found that the prefrontal cortex
tends to get a little bit smaller as we get older. If the prefrontal
cortex is shrinking as we get older and working memory depends
on the prefrontal cortex, then we can understand why working
memory might tend to get worse as we age.
QUESTIONS TO CONSIDER
2 Considering the acronym EARS, are there any changes you’d like
to make in your life that might help you age more gracefully?
Dyslexia
•• Perhaps the most common SLD, and also the best known, is
dyslexia. In fact, some reports estimate that as many as 80% of
children with a learning disability suffer from dyslexia. Dyslexia
has also received more attention from scientists than any other
learning disability, and a great deal has been learned about what’s
going on.
•• When you’re first starting to read, written words are just unfamiliar
strings of letters, and you have to go through a sequence of steps
to figure out what word those letters correspond to. And a critical
step in that process is breaking down the sound of the word into
its component phonemes.
•• Both of these brain regions are activated during reading, too, even
though you’re not normally hearing or speaking when you read.
But in addition to these regions, reading also activates regions in
the visual cortex, such as the visual word form area, which is on
the bottom surface of the left hemisphere of brain, toward the
back. It’s critically important in the visual recognition of letters
and written words.
QUESTIONS TO CONSIDER
Learning has a lot more to do with what the student does than
with what the teacher does. As the great psychologist Herbert
Simon put it, “Learning results from what the student does and
thinks, and only from what the student does and thinks. The
teacher can advance learning only by influencing what the student
does to learn.”
•• And this is how the most effective learners keep learning. They
don’t let obstacles get in the way; instead, they find ways to
overcome difficulties and continue their learning.
•• But it’s not just the work world where you can see the advantages
of active learning. Imagine trying to learn algebra without doing
any practice problems. It just wouldn’t work. You can read the
textbook over and over, but until you’ve solved a bunch of actual
problems, you’ll never master algebra.
•• How can we be more active in our learning so that we can get the
most out of it? One idea is to intentionally ask questions while
we’re learning. Suppose that you want to learn about gardening,
so you’re reading a book on the topic. One excellent way to get
the most out of your reading is to try to come up with a set of key
questions that the book answers.
•• But where are you going to get the test? You’ll occasionally
run across a book that includes some questions, but that’s the
exception, not the rule. But if you’ve adopted the previous
suggestion and made up a quiz on the material you’re trying to
learn, then you’re all set! Just wait until the next day or week and
then try taking the quiz you made up. The exercise of taking that
test will help consolidate what you’ve learned. And if there are
questions you can’t answer, then that tells you exactly what you
might want to review.
•• A final practical way to make your learning more active and less
passive is to try to teach others what you’re learning. There’s no
better way to expose holes in your understanding than to try to
explain something to other people.
•• The same goes for the neural network inside your skull. If you train
your brain on information that is unreliable or biased, it will learn
generalizations that may be unreliable or flat-out wrong.
•• So, if you want to optimize your learning, then it’s crucial to give
your neural network information that is as reliable and unbiased
as possible. But how can we do that?
•• There are many things we can do to keep our brains healthy, but
let’s focus on 3 that scientific studies have identified as particularly
important: staying active, eating right, and getting enough sleep.
•• Study after study has found that your diet not only influences
your health and longevity, but it also can have a dramatic impact
on your mental functions. For example, eating a relatively low-
calorie Mediterranean diet has not only been found to improve
heart health, but also to improve cognitive function and even
significantly reduce the risk of Alzheimer’s disease.
SUGGESTED READING
Duckworth, Grit.
Dweck, Mindset.
Ericsson and Pool, Peak.
Ratey and Hagerman, Spark.
Waldrop, “The Science of Teaching Science.”
2 Are there any strategies that weren’t in this lecture but that you’ve
found helpful in optimizing your learning and memory?
Buettner, Dan. The Blue Zone: Lessons for Living Longer from the People
Who’ve Lived the Longest. Washington, DC: National Geographic,
2008.
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