232 Chapter 6 MeMory

8. Explain why recognition tests of memory generally produce
better results than recall tests.
(e) _______ tasks (such as multiple-choice questions) gen-
erally produce better memory retrieval than (f) _______
tests (free recall, serial recall, or paired-associates recall)
because they provide more retrieval cues that help jog
memory.
9. Describe the causes of amnesia and the two major types
of amnesia.
Amnesia, or memory loss, may be caused by psychological
factors or by physical factors such as degenerative brain dis-
eases and brain trauma. There are two general types of am-
nesia: (g) _______ amnesia (loss of memory for past events)
and (h) _______ amnesia (inability to form new memories).

Recall It
1. What type of interference accounts for why you may forget to 3. When it comes to remembering what you’ve learned, _______
advance the year when writing checks early in a new year? practice is preferable to _______ practice.

2. Which of the following is not a helpful way to reduce the 4. Memory loss in which earlier life events are forgotten is
effects of interference on memory? known as
a. Avoid overlearning. a. dissociative amnesia.
b. Study material just before going to bed. b. retrograde amnesia.
c. Rehearse or practice material repeatedly. c. retroactive amnesia.
d. Avoid studying similar content simultaneously. d. anterograde amnesia.

Think About It
■ Have you had any tip-of-the-tongue experiences? Were ■ Why is it not a good idea to apply the principle of
you eventually able to retrieve the memory you were “massed practice” when preparing for exams? What
searching for? If so, how were you able to retrieve it? study techniques are likely to be more effective?

Recite It answers placed at the end of chapter.

MODULE 6.3 The Biology of Memory

10 Identify the key brain structures involved in memory and explain the roles of
neuronal networks and long-term potentiation.
11 Explain the role that genetics plays in memory.
12 Apply knowledge of how memory works to power up your memory.

How are memories formed in the brain? Where are they stored? Breakthrough
research is beginning to answer these and other questions that probe the biological
underpinnings of memory. In this module, we examine what is presently known
about those underpinnings.

Brain Structures in Memory:

engram Lashley’s term for the physical
trace or etching of a memory in the brain.
Where Do Memories Reside?
Psychologist Karl Lashley (1890–1958) spent much of his career attempting to track
down the elusive engram, the term he used to describe a physical trace or etching in
the brain where he believed a memory is stored. A rat that learns to run a maze, for
example, should have an engram somewhere in its brain containing a memory trace
of the correct route leading to the exit or goal box.
Lashley spent years training rats to run mazes, then surgically removing parts
of their cerebral cortexes, and testing them again to see if their memories for mazes

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 M O D U L E 6.3 233

remained intact. He reasoned that if removal of a part of the cortex wiped away a CONCept 6.23
given memory, that part must be where the particular memory was stored. Despite Memories are stored in complex networks
years of painstaking research, he found that rats continued to run mazes they had of interconnected brain cells called neu-
learned previously regardless of the parts of the cortex he removed. The rats simply ronal networks.
did not forget. He concluded that memories are not housed in any specific brain
structure but must be scattered about the brain.

Neuronal Networks: The Circuitry of Memory

Memory scientists today believe that memories are not etched into particular neurons in
the brain but rather are encoded and stored in memory circuits that are housed through-
out the brain in complex networks of neurons called neuronal networks (also called neu-
ral networks) (Lu & Zuo, 2015; Miller, 2015). In simplest terms, think of
these memory networks as intricate webs of connections among neurons in
the brain. We just don’t know how many neurons are involved in storing any
individual memory—perhaps only few thousand or fewer, or perhaps even
hundreds of millions of neurons dispersed in complex networks throughout
the brain (Quiroga, Fried, & Koch, 2013).

The Hippocampus: A Storage Bin for Memory

The hippocampus, a seahorse-shaped structure in the forebrain, is essential
to forming memories of facts and general information (semantic memory)
and life experiences (episodic memory) (Buzsáki, 2013; Merkow, Burke,

Roger Harris/Science Source

& Kahana, 2015).
The hippocampus doesn’t seem to be involved in procedural memory,
the kind of memory process we draw upon when riding a bicycle or using
tools. Nor does the hippocampus appear to be the final destination for new
memories. Rather, it appears to be a temporary storage bin for holding new
memories, perhaps for a few weeks or months, before they are moved and
filed away in the cerebral cortex and other parts of the brain for long-term storage
(Aamodt & Wang, 2008; Stickgold & Wehrwein, 2009).
If you suffered extensive damage to your hippocampus, you might develop antero-
CONCept 6.24
grade amnesia and be unable to form new memories. Depending on the extent of the
Damage to the hippocampus could
damage, you might retain earlier memories but each new experience would fail to leave prevent you from forming new memories,
any mark in your memory (Thompson, 2005). so that you might be unable to remember
Memory also depends on other brain structures, including the thalamus and the someone you’ve just met.
amygdala. Damage to the thalamus can result in amnesia. The amygdala plays an im-
portant part in encoding emotional experiences, such as fear and anger. Scientists believe CONCEPT LINK
the amygdala and hippocampus are especially active during emotionally charged experi- The hippocampus plays a role in
ences, which serves to strengthen and preserve memories of such meaningful life events remembering the context in which
(Hassert, Miyashita, & Williams, 2004). All in all, there is no one memory center in the fearful responses were experienced. See
brain, no one part of the brain entirely responsible for memory formation. Module 8.4.

Strengthening Connections Between Neurons:

The Key to forming Memories
Locating neuronal networks corresponding to particular memories makes finding the
proverbial needle in the haystack seem like child’s play. The human brain contains bil-
lions of neurons and trillions of synapses among them. Individual neurons in the brain
can have thousands of synaptic connections with other neurons. In tracking down spe-
cific networks of cells where memories are formed, researchers turned to a relatively
simple animal, a large sea snail (Aplysia) that possesses a mere 20,000 neurons. neuronal networks Memory circuits
The landmark research that Eric Kandel, a molecular biologist and Nobel Prize in the brain that consist of complicated
winner, performed on Aplysia represented a major step forward in unraveling the networks of nerve cells.

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234 Chapter 6 MeMory
Eric R. Kandel
Nobel Prize winner Erik Kandel holding
an Aplysia, the sea snail he used to study
the biological bases of memory.
CONCept 6.25
The key to forming memories may lie
in strengthening the interconnections
between the neurons that form neuronal
networks in the brain.
CONCept 6.26
Scientists suspect that long-term potenti-
ation (LTP) may be needed for long-term
memory to occur.
CONCept 6.27
Scientists have begun to unravel the genetic
bases of memory, which may lead to the
development of safe drugs that can help
preserve or restore memory functioning.
long-term potentiation (LTP) The long-
term strengthening of neural connections
as the result of repeated stimulation.
Copyright 2018 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-203
biological bases of memory (Kandel, 1995; Kandel & Hawkins, 1993). Because learn-
ing results in the formation of new memories, Kandel needed to first demonstrate that
these animals were capable of learning new responses. To accomplish this, he and his
colleagues first desensitized the snails to receiving a mild squirt of water. After a num-
ber of trials, the animals became habituated to the water squirt so that it no longer
caused them to budge. In the second phase of the experiment, the researchers paired
the squirt with a mild electric shock. The animals showed they could learn a simple
conditioned response—reflexively withdrawing their gills (their breathing apparatus)
when squirted with water alone. This self-defensive maneuver is the equivalent of the
snails’ battening down the hatches in anticipation of impending shock (Rupp, 1998).
Kandel observed that the amount of neurotransmitters released into synapses
between the nerve cells that control the withdrawal reflex increased as the animals
learned the conditioned response. The added neurotransmitter kicked the reflex into
overdrive, making it more likely to fire. In effect, these synapses became stronger—
that is, more capable of transmitting neural messages. Kandel had shown that mem-
ory formation involves biochemical changes occurring at the synaptic level.
“Cells That Fire Together, Wire Together”
Synaptic connections between neurons can be strengthened by repeated electrical
stimulation. In effect, neurons that “fire together, wire together.” A long-lasting in-
crease in the strength of synaptic connections is called long-term potentiation (LTP).
The word potentiation simply means strengthening. When synaptic connections be-
come stronger, neurons are better able to communicate with each other (Monaco
et al., 2014; Sejnowski & Delbruck, 2012).
The ability to form long-term memories appears to depend on strengthening synaptic
connections between neurons within complex brain circuits (Craddock, Tuszynski, &
Hameroff, 2012). In other words, we remember life experiences or newly learned infor-
mation (such as the meaning of the word potentiation) when groups of neurons
in the brain form connections in which the particular memory is embedded or
encoded (Tayler et al., 2012). We can think of a memory as a pattern of neural
activity within a particular brain circuit. Retrieving a particular memory from
storage (“Let’s see, what does potentiation mean?”) brings the information to
mind by reactivating the corresponding pattern of brain activity.
Scientists believe the process of LTP plays a key role in converting
short-term memory into long-term memory (Hutchison, Chidiac, &
Leung, 2009). But just how are synaptic connections strengthened? One
Artpartner-images/The Image Bank/Getty Images
way is to repeatedly practice or rehearse information you want to remem-
ber (studying, anyone?). However, mere repeated practice or rote memo-
rization may not be sufficient to remember complex material, such as the
principles of memory discussed in this chapter. We may need to strengthen
newly formed memories by elaborating on the meaning of the material—
for example, by thinking of examples of the concepts discussed in this
chapter or by relating them to your own personal experiences.
Genetic Bases of Memory
Promising research with genetic engineering is offering new insights into how mem-
ory works. Transforming short-term memory into long-term memory depends on
the production of certain proteins that are regulated by certain genes (Ramamoor-
thi et al., 2011). Learning more about how these genes functions will help unravel
the molecular bases of memory, perhaps leading to a time when we will be able to
directly manipulate “memory genes” to enhance learning and memory ability. Sci-
entists have already been able to boost memory ability in mice by means of genetic
 m o d u l e 6.3 235

Concept Chart 6.3 Biology of memory: Key Concepts

Concept description

Lashley’s engram Despite years of research, Karl Lashley failed to find

evidence of an engram, his term for a physical trace or

©Jurgen Ziewe/Shutterstock.com
etching in the brain where he believed a memory is stored.

Neuronal networks Memory scientists believe that memories may “reside” in

complex networks of neurons distributed across different
parts of the brain.

Biological underpinnings The hippocampus is a key brain structure in converting

of memory short-term memory into long-term memory. The
conversion of short-term to long-term memory appears
to depend on long-term potentiation, the process of
strengthening synaptic connections between neurons
in which memories are embedded.

Genetic factors in memory Conversion of short-term memory to long-term memory

depends on brain proteins whose production is regulated
by certain genes. Advances in genetic engineering show
that it is possible to enhance learning and memory ability
in nonhuman organisms by genetic manipulation.

engineering (tinkering with genes) (Zhu et al., 2011). Further advances in genetic en-
gineering might also one day reap benefits in helping people suffering from memory
loss, such as those with Alzheimer’s disease.
Scientists hope that knowledge gained about the role of brain proteins in memory
and the genes that help regulate their production may eventually lead to the devel-
opment of drugs to treat or even cure Alzheimer’s disease and other memory disor-
ders. Perhaps we’ll even have drugs that boost the memory functioning of normal
individuals. In the meantime, think critically if you encounter claims about so-called
memory-enhancing drugs. We have no compelling scientific evidence that any drug or
supplement available today enhances memory in normal individuals (Begley, 2011a).
Concept Chart 6.3 summarizes some of the key concepts relating to the biology
of memory.

Applying psyChology
Powering Up Your Memory
Even if you never compete in a memory championship, you can learn to boost your CONCEPT 6.28
memory power. Techniques specifically aimed at enhancing memory are called You can boost your memory power in
mnemonics, some of which have been practiced since the time of the ancient Greeks. many ways, such as by using mnemon-
Yet perhaps the most important ways to power up your memory are to take care of ics, focusing your attention, practicing
your health and to adopt more effective methods of studying, such as the SQ3R1 repeatedly, taking care of your health, and
system (see “A Message to Students” in the preface of the text). adopting effective study habits.

using mnemonics to Improve memory

A mnemonic is a device for improving memory. The word mnemonic is derived from the
name of the Greek goddess of memory, Mnemosyne, and is pronounced neh-MAHN-ik mnemonic A device for improving
(the first m is silent). Here are some of the most widely used mnemonic devices. memory.

Copyright 2018 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-203