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Annual Review of Psychology Vol 56 2005 1st Edition
Susan T. Fiske Digital Instant Download
Author(s): Susan T. Fiske, Daniel L. Schacter, Alan E. Kazdin
ISBN(s): 9780824302566, 0824302567
Edition: 1
File Details: PDF, 4.24 MB
Year: 2005
Language: english
Annu. Rev. Psychol. 2005.56:1-23. Downloaded from arjournals.annualreviews.org
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December 8, 2004
12:13
Annual Reviews
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10.1146/annurev.psych.56.091103.070239

Annu. Rev. Psychol. 2005. 56:1–23

doi: 10.1146/annurev.psych.56.091103.070239
Copyright
c 2005 by Annual Reviews. All rights reserved
First published online as a Review in Advance on June 10, 2004

IN SEARCH OF MEMORY TRACES

Richard F. Thompson
Neuroscience Program, University of Southern California, Los Angeles,
California 90089-2520; email: [email protected]
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Key Words learning, hippocampus, cerebellum, inactivation, localization

■ Abstract The key issue in analyzing brain substrates of memory is the nature of
memory traces, how memories are formed, stored, and retrieved in the brain. In order
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to analyze mechanisms of memory formation it is first necessary to find the loci of

memory storage, the classic problem of localization. Various approaches to this issue
are reviewed. A particular strategy is proposed that involves a number of different
techniques (electrophysiological recording, lesions, electrical stimulation, pathway
tracing) to identify the essential memory trace circuit for a given form of learning and
memory. The methods of reversible inactivation can be used to localize the memory
traces within this circuit. Using classical conditioning of eye blink and other discrete
responses as a model system, the essential memory trace circuit is identified, the basic
memory trace is localized (to the cerebellum), and putative higher-order memory traces
are characterized in the hippocampus.

CONTENTS
INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Simplified Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Strategies to Study Memory Formation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
OUR MODEL SYSTEM: EYEBLINK CONDITIONING . . . . . . . . . . . . . . . . . . . . . 6
Motor Control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
The Hippocampus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
The Essential Circuitry: The Cerebellum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
CONCLUSION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

INTRODUCTION
A major achievement of recent research on brain substrates of learning and mem-
ory, to which our work has contributed, is the recognition that there are several
different forms or aspects of memory involving different brain systems (Figure 1).
Squire (1992) has argued eloquently for the distinction between declarative and
nondeclarative forms of memory, as has Schacter (1987). The distinction be-
tween episodic and semantic memory—“What did you have for breakfast?” versus
“Where is the Eiffel Tower?”—has been stressed by Tulving (1985).
0066-4308/05/0203-0001$14.00 1
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Figure 1 A current view of the various forms of learning and memory and their putative brain substrates.
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LOCALIZATION OF MEMORY TRACES 3

Implicit or nondeclarative memory is very much a grab bag. In general, explicit

memory involves awareness of the memory whereas implicit memory does not
necessarily involve being aware of the memory (aware meaning verbal report).
The schema of Figure 1 is of course oversimplified. When an organism learns
something, a number of brain systems can become engaged. However, in most
cases there is one critical brain system, which when damaged causes permanent
impairment in the particular form of learning and memory.
Many readers will recall Lashley’s (1950) pessimistic conclusion that his series
of experiments “has yielded a good bit of information about what and where
the memory trace is not” in his famous article, “In Search of the Engram.” In
all fairness to Lashley, the existence of several different forms of memory with
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differing neuronal substrates was not recognized at that time, nor were modern
analytic techniques available then.
We adopt the following definitions from an earlier review (Lavond et al. 1993).
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The essential memory trace, together with its associated circuitry, is necessary and
sufficient for the basic aspects of a given form of learning (e.g., acquisition and
retention). Other brain structures may also form memory traces, defined loosely as
learning-induced changes in neuronal activity, but these may not be essential for
the basic learning. Thus, in eyeblink conditioning, long-lasting learning-induced
increases in neuronal activity develop in the hippocampus (Berger et al. 1976), but
the hippocampus is not necessary for basic delay associative learning and memory
(Schmaltz & Theios 1972).
Aversive classical conditioning can modify the receptive-field properties of neu-
rons in sensory systems. Particularly striking are the studies of Weinberger and
associates (Edeline & Weinberger 1991, Weinberger et al. 1984) showing that neu-
rons in secondary areas of auditory thalamus and cortex shift their best frequencies
toward the frequency of the conditioning stimulus in aversive Pavlovian condition-
ing (see also Bao et al. 2003). The motor area of the cerebral cortex provides yet
another example. Classical eyeblink conditioning results in marked and persist-
ing increases in excitability of pyramidal neurons in motor cortex (Woody et al.
1984). However, the motor cortex is not necessary for either learning or memory of
the conditioned eyeblink response (Ivkovich & Thompson 1997). These “higher-
order” memory traces appear to play important roles in the adaptive behavior of
the organism.
The obvious point here is that the organism can learn, remember, and per-
form the basic learned response following destruction of nonessential memory
trace systems, but destruction of the essential memory trace system abolishes this
ability. We draw a distinction between the essential memory trace and the es-
sential memory trace circuit. The latter includes the necessary input and output
circuits as well as the essential memory trace itself. These are all identified by
lesions. But lesions, per se, cannot distinguish between the essential trace and the
essential circuitry; lesions of either completely prevent and abolish the learned
response.
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4 THOMPSON

Simplified Systems
How does one go about finding memory traces? One very productive approach is
the use of simplified “model” systems, emphasized early by Thompson & Spencer
(1966) (see also Kandel & Spencer 1968). Thus, vertebrate spinal reflexes and the
gill-withdrawal reflex circuit in Aplysia have both served as very productive models
to analyze processes of habituation and sensitization. In the case of habituation,
at least, the memory trace is embedded within the reflex pathway under study so
lesions abolish the reflex as well as the trace. For monosynaptic pathways in both
Aplysia and spinal cord the mechanism of short-term habituation appears to be
simply a decrease in the probability of transmitter release as a result of repeated
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activation, a presynaptic process of synaptic depression (Farel & Thompson 1976,

Kandel 1975).
But even for a simple behavioral phenomenon like habituation, the neural sub-
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strate can prove to be complex, particularly for long-lasting habituation, e.g., be-
tween sessions as opposed to within session response decrements. In the Aplysia
cellular monosynaptic system, Ezzeddine & Glanzman (2003) have shown that
prolonged habituation depends on protein synthesis, protein phosphatase activ-
ity, and postsynaptic glutamate receptors. Indeed, their data indicate that stimuli
inducing habituation of the gill-withdrawal reflex appear to activate sensitizing
processes as well, as proposed in the “dual-process” theory of habituation by
Groves & Thompson (1970).
Thus, much is still not known about the detailed mechanisms of memory traces
in this simplest form of learning. By mechanisms we mean the physical-chemical
substrate of memory store. Although there are many candidate mechanisms, as of
this writing we do not yet know the detailed physical bases of memory storage for
any form of learning and memory in the mammalian brain. But understanding the
mechanisms of memory storage will not inform us of what the memories are. The
actual memories are coded by the neuronal circuits that code, store, and retrieve
the memories, what I term here the “essential circuits.”

Strategies to Study Memory Formation

There appear to be two general strategies for the study of memory storage. One
approach is to pick a mechanism of neural plasticity such as long-term potentiation
(LTP) in a structure known to be important for memory and attempt to show that it
is a substrate for memories. Richard Morris and associates published a heroic effort
to demonstrate that activity-dependent synaptic plasticity, particularly LTP (and
LTD, long-term depression), “is both necessary and sufficient for the information
storage underlying the type of memory mediated by the brain area in which that
plasticity is observed” (Martin et al. 2000). They established several “formal” cri-
teria by which to judge this hypothesis. We examine these criteria below but here
we simply note their conclusion that “synaptic plasticity is necessary for learn-
ing and memory, but that little data currently supports the notion of sufficiency”
(p. 649).
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LOCALIZATION OF MEMORY TRACES 5

Actually, there are many forms of plasticity in the nervous system, synap-
tic plasticity being only one. There are also changes in neuron excitability due
to factors intrinsic to the neurons, as in decreases in after-hyperpolarization, as
well as structural changes in dendrites, formation of new neurons, and actions
of glial cells. LTP has been the most intensely studied putative synaptic mech-
anism of memory storage, particularly in the hippocampus (e.g., Martin et al.
2000), but the evidence is still far from clear. There is a vast literature analyz-
ing mechanisms of LTP but very little work attempting to show that it is a basis
for the storage of memories (see Shors & Matzel 1997, Wilson & Tonegawa
1997).
The alternative strategy is to begin with a well-characterized form of learning
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and memory and determine the memory traces involved. In order to do this it is first
necessary to find where in the brain the memories are stored, the classical problem
of localization. Because learning involves changes in behavior as a result of expo-
by Ball State University on 01/05/09. For personal use only.

sure to stimuli that do not change, there must be alterations at some loci in the brain.
A given form of learning may involve one or multiple loci of memory storage, but
they must exist. We have used this strategy in our search for memory traces.
How does one go about finding a memory trace in the behaving mammal? Olds,
Disterhoft, and associates wrestled with this issue in pioneering studies (Olds et al.
1972). Their basic approach was to record extracellular unit cluster discharges in
many brain areas in freely moving rats, looking for increases in unit response on
the conditioning day that were significantly greater than responses to the stimuli
on the preceding pseudoconditioning day, i.e., a learning-induced increase in unit
activity. The behavioral situation was simply an auditory conditioned stimulus
(CS+) followed after 2 sec by a food pellet unconditioned stimulus (US) versus a
CS− with no food pellet. The studies were very well controlled.
The initial criterion they set for identification of the memory trace was shortest
latency learned responses:

The “earliest” conditioned brain responses (i.e., those appearing with the
shortest latencies after application of the CS) might thus be considered to be
“at the site” of conditioning, and other later conditioned brain responses might
be considered to be secondary to them. Similarly (and hopefully validating
this supposition) “early” new brain responses might be expected to emerge at
the site of old responses indicating this site to be a junction point between old
and new (Olds et al. 1972, p. 202).

In a most interesting thought piece, Michael Gabriel (1976) argued against the
use of the shortest latency response as the criterion for identification of the memory
trace. He proposed an alternative “bias” hypothesis. In essence, he suggested that a
tonic flow of impulses originating from a distant neuronal locus biased the activity
of the neurons being recorded as the shortest latency learned response. So the
increase of the shortest latency unit response could be due to tonic actions on
these neurons from a distant site where learning has resulted in a tonic increase
in activity. Gabriel draws the following rather strong conclusion: “clearly the bias
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6 THOMPSON

hypothesis argues against the idea of mapping the brain for short-latency neuronal
responses in order to localize engrams” (p. 279).
In a companion piece to the Olds et al. (1972) article, Disterhoft & Olds (1972)
do consider the possibility of tonic influences, as Gabriel notes. They suggest an
additional criterion, namely the earliest increase in neuronal activity that develops
over the trials of training. I return to this issue below.
In the late 1960s and early 1970s we tried several different types of model
systems to analyze neural substrates of mammalian associative learning and mem-
ory. Because of our success in using spinal reflexes for analysis of habituation
we tackled classical conditioning of the flexion reflex in the acute spinal animal,
with some success (Patterson et al. 1973). However, I concluded it was not a good
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model of associative learning in the behaving mammal.

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OUR MODEL SYSTEM: EYEBLINK CONDITIONING

Thanks in part to efforts by Michael Patterson, then a postdoc in my lab who had
obtained his Ph.D. with Isadore Gormezano, and because of the elegant and com-
prehensive behavioral studies by Gormezano, we adopted his preparation: classical
conditioning of the nictitating membrane (NM) eyeblink response in the restrained
rabbit. We have detailed the advantages of this preparation elsewhere (Thompson
et al. 1972, Thompson et al. 1976). Perhaps the most important advantages are:
(a) The unconditioned response (UR) provides an independent measure of per-
formance against which to compare effects of variables acting on the conditioned
response (CR), and (b) the behavioral conditioned response is robust, reliable, and
discrete, and the exact amplitude-time course of the response is measurable. There
were a number of other advantages as well: no sensitization, pseudoconditioning,
or alpha responding; many trials required to learn; and extensive parometric data on
the properties of the CR (see Gormezano et al. 1983, and the important theoretical
analyses by Wagner and associates, e.g., Wagner & Donegan 1989).

Motor Control
We began our analysis with a focus on the motor nerves and nuclei that gener-
ated the reflex and learned eyeblink responses (Cegavske et al. 1976, Cegavske
et al. 1979, Young et al. 1976). To oversimplify from our work and the work of
others, the sixth nerve was critically important for the nictitating membrane (NM)
response, but the third and fourth nerves were also involved and the seventh nerve
was critical for closure of the external eyelid (obicularis oculi muscles). In terms
of motor nuclei, the seventh nucleus controlled external eyelid closure and the
accessory portion of the sixth nucleus innervated the retractor bulbous muscle,
which retracted the eyeball and forced the nictitating membrane out across the
front surface of the eye. The fourth and sixth nuclei acted synergistically with
the accessory sixth and seventh. We showed with simultaneous recordings that
NM extension and external eyelid closure [actually electromyographic (EMG)
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LOCALIZATION OF MEMORY TRACES 7

activity of the obicularis oculi muscles] were essentially perfectly correlated over
the course of learning (McCormick et al. 1982b). Indeed, obicularis oculi EMG
appears to be the most robust and sensitive measure of learning (Lavond et al.
1990). Recordings from the relevant motor nuclei, particularly the sixth, acces-
sory sixth, and seventh, showed essentially identical patterns of learning-induced
increases in neuronal activity.
It is important to stress the fact that the pathways mediating the reflex eye-
blink are in the brain stem and do not involve “higher” brain systems such as the
cerebellum and hippocampus. Specifically, there are direct projections from the
trigeminal nucleus (activated by stimulation of the cornea and surrounding tissues)
to the relevant motor nuclei and indirect projections relaying via the brain stem
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reticular system to the motor nuclei (Hiraoka & Shimamura 1977).

The basic logic of our approach seemed reasonable—identify the critical motor
neurons involved in control of the learned response and trace the essential circuitry
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backward to the source, the “engram.” The fact that several motor nuclei showed the
same pattern of learning-induced activation argued against any plasticity unique
to one motor nucleus (e.g., the accessory sixth) or to processes of plasticity at
the level of the motor nuclei. Instead, it seemed most likely that the learning-
induced response in the motor nuclei was driven from a common central source.
The pattern of learning-induced increase in the activity of neurons in the motor
nuclei in the CS period in paired trials and on CS-alone trials (i.e., the neuronal
CR) was strikingly similar in form to the amplitude-time course of NM extension,
external eyelid closure, and of course the EMG recorded from the obicularis oculi
muscles. Indeed, an envelope of increased frequency of discharge of neurons in
the motor nuclei preceded in time and closely predicted the form of the behavioral
eyeblink-NM response (Figure 2). This close predictive parallel was most evident
with unit cluster recordings but was also true for single unit recordings in the motor
nuclei.

USE OF THE MOTOR NEURON NEURONAL MODEL Our initial logic—to work back-
ward from the motor nuclei—was not as simple as it might have seemed because of
the very large number of central brain systems that project to the motor and premo-
tor nuclei. So we adopted a different strategy. It was apparent that the amplitude-
time course form of the conditioned eyeblink-NM response closely paralleled and
followed the pattern of increased unit activity in the motor nuclei. So we focused
on this behavioral model of the learning-induced increase in neural activity in
the motor nuclei and used it as a template or model (Figure 2). The higher brain
systems that acted to generate and drive the learning-induced neuronal CR, the
“model” in the motor nuclei, must show the same pattern of learning-induced ac-
tivity as do the motor nuclei, and hence the amplitude-time course of the learned
eyeblink-NM response. Thus, we searched through higher brain systems looking
for learning-induced neuronal activity that correlated closely with and preceded in
time the form of the learned eyeblink-NM response. In essence, we mapped vir-
tually the entire brain of the rabbit in 1 mm steps, searching for neuronal models
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8 THOMPSON
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Figure 2 Histograms of unit cluster recordings from the ABD and simultaneous recording of
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the eyeblink response (NM, the “third eyelid”) in classical conditioning in the rabbit summed
and averaged over eight trials before (A) and after (B) learning. First cursor onset of tone CS,
second cursor onset of corneal airpuff US, trace duration 750 msec, upward movement of
the NM trace indicates extension (eyelid closure). Note that the pattern of neural discharges
precedes and models the amplitude-time course of the behavioral response. ABD, abducens
motor nucleaus; CS, conditioned stimulus; NM, nictitating membrane; US, unconditioned
response. (From Cegavske et al. 1979.)

of the learned behavioral response (see, e.g., McCormick et al. 1983, Thompson
et al. 1976). We did not search blindly but rather system-by-system.
The basic assumption is that the neuronal sites of memory trace formation will
show a pattern of increased frequency of unit cluster and single-unit discharges
that precedes and closely predicts the pattern of response in the motor nuclei and
hence in the amplitude-time course of the behavioral CR. We felt this criterion
was preferable to the earlier ideas of shortest latency response, earliest response
over trials, or first site to show increased tonic responses. This assumption proved
to be the key that permitted us to localize a memory trace.

The Hippocampus
Because of its involvement in memory, we began with the hippocampus (Berger
et al. 1976). To our delight, both unit cluster recordings and single-unit responses
recorded from CA1 and CA3 showed the requisite learning-induced responses, i.e.,
pattern of increased discharge frequency that preceded in time (within trials) and
correlated virtually perfectly with the amplitude-time course of the behavioral CR
(Figure 3).
We showed that this neuronal model of a memory is generated by identified
pyramidal neurons (Berger & Thompson 1978b). Furthermore, the hippocampal
unit response began to develop in the US period within just a few trials of training
(a criterion by Olds et al. 1972) and moved into the CS period in close association
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LOCALIZATION OF MEMORY TRACES 9

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Figure 3 Behavioral eyeblink (nictitating membrane) and hippocampal unit response

averaged over 100 trials in a well-trained rabbit. The unit response is of a single, iden-
tified pyramidal neuron in CA1. Upper trace, raw single 100-msec sweep to show the
neuron response, middle trace averaged NM response, lower trace cumulated histogram
of unit activity; cursors and durations in middle and lower traces as in Figure 2. Note
that the pattern of response of this neuron both precedes and models the amplitude-time
course of the behavioral response. (From Berger & Thompson 1978b.)

with the development of the behavioral CR (Berger & Thompson 1978a). It seemed
like a perfect candidate for the engram. In addition, electroencephalogram (EEG)
activity recorded from the hippocampus at the beginning of training predicted the
rate of learning (Berry & Thompson 1978). We characterized this learning-induced
response in the hippocampus in some detail (see Berger et al. 1986). Unfortunately,
as we noted above, animals could learn the basic delay-conditioned NM-eyeblink
response following hippocampal lesions (Schmaltz & Theios 1972).
This apparent enigma illustrates a fundamental limitation of using neuronal
recordings to localize memory traces. The fact that a neuronal model of the learned
response occurs at a given locus does not necessarily mean it originates there.
Indeed, the learning-induced neuronal model in the hippocampus, at least in the CS
period, is abolished by appropriate cerebellum lesions (Clark et al. 1984). Actually,
a number of loci in the brain exhibit the learning-induced neuronal model of the
learned behavioral response. I return to this issue below.
The breakthrough insofar as the hippocampus is concerned came in a study in
our laboratory by Paul Solomon and Donald Weisz using the trace-conditioning
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10 THOMPSON

paradigm, where a 500-msec period of no stimuli separated CS offset from US

onset (Solomon et al. 1986). In brief, large bilateral hippocampal lesions made
before training markedly impaired subsequent learning of the trace (but not delay)
eyeblink CR, a result replicated exactly by Moyer et al. (1990). If the hippocampal
lesions are made immediately after learning, the trace CR is abolished (with no
effect on the delay CR), but if the lesions are made a month after training the trace
CR remains intact (Kim et al. 1995).
In sum, hippocampal lesions induced anterograde amnesia (impairment of
postlesion learning) and marked but time-limited retrograde amnesia for the trace
eyeblink CR. These are precisely the effects that such lesions have on declarative
memory in humans (Squire 1987). These results suggest that trace eyeblink condi-
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tioning provides a simple model of hippocampal-dependent declarative memory,

a possibility strongly supported by studies of humans with hippocampal-medial
temporal lobe amnesia. Such patients are markedly impaired on acquisition of trace
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eyeblink conditioning if the trace interval is sufficiently long but not impaired at
all on delay conditioning (see McGlinchey-Berroth et al. 1997).
Clark & Squire (1998, 1999) made the striking observation that awareness of
the training contingencies in normal human subjects correlated highly with the
degree of learning of trace eyeblink conditioning. Recall that awareness is a key
defining property of declarative memory. They also showed that awareness played
no role in delay conditioning, and that expectancy of US occurrence influenced
trace but not delay conditioning (Clark & Squire 2000, Clark et al. 2001). They
conclude that delay and trace conditioning are fundamentally different phenomena,
with delay conditioning inducing nondeclarative or procedural memory and trace
conditioning inducing declarative memory. Hence, trace eyeblink conditioning
in rabbits would seem to provide a very elementary instantiation of declarative
memory in animals.
These strikingly parallel results of hippocampal lesions in rabbits and humans
for trace eyeblink conditioning suggest the possibility that a memory trace(s) may
develop in the hippocampus in trace conditioning, but do not prove it. Recall that
delay eyeblink conditioning results in the development of a pronounced neuronal
model of the learned response in the hippocampus (as does trace training).
Several lines of evidence support the view that localized changes do develop
in the hippocampus in eyeblink conditioning. Thus, the properties of increased
neuronal activity are strikingly similar to the properties of LTP (Berger et al. 1986,
Thompson 1997, Weisz et al. 1984), and induction of LTP in the hippocampus
can enhance eyeblink learning (Berger 1984). There is also a dramatic learning-
induced decrease in the after-hyperpolarization in pyramidal neurons in eyeblink
conditioning, most evident in trace conditioning, which leads to increased ex-
citability. This alteration is intrinsic to the neurons due to a decrease in one or more
calcium-dependent outward potassium currents (Disterhoft et al. 1986, Disterhoft
& McEchron 2000). There is also a dramatic increase in dendritic membrane-
associated protein kinase C after eyeblink conditioning (Olds et al. 1989). After
trace eyeblink conditioning there is a substantial increase in the number of multiple
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LOCALIZATION OF MEMORY TRACES 11

synapse boutons in hippocampal neurons (Geinisman et al. 2001). Finally, there

is a dramatic increase in the number of new neurons in the hippocampus in trace,
but not delay, eyeblink conditioning (Gould et al. 1999, Shors et al. 2001).
These sets of structural and chemical changes are certainly consistent with the
formation of higher-order memory traces in the hippocampus. But recall that cere-
bellar lesions can abolish both the behavioral learned response and the increased
neuronal response in the CS period in the hippocampus in both delay and trace
learning (see below and Clark et al. 1984, Woodruff-Pak et al. 1985). So doubts
remain. Interestingly, the early (over trials)-developing increase in neuronal activ-
ity in the hippocampus in the US period is not completely abolished by cerebellar
lesions (Clark et al. 1984). In my view, it is necessary to identify the entire essential
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circuitry from stimulus input to motor output in order to establish definitively that
the essential memory trace, e.g., for trace conditioning, develops and is stored,
albeit temporarily, in the hippocampus (see below). To date, not all the essential
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circuitry has been identified (see Thompson & Kim 1996). Yet another remaining
puzzle is where the trace memories go after the hippocampus is no longer necessary.

The Essential Circuitry: The Cerebellum

So the hippocampus is not the locus of the essential memory trace for standard-
delay classical eyeblink conditioning. In a detailed mapping study of the brain stem
we identified several loci where neurons exhibited the requisite neuronal model
of the CR—increase in frequency of discharge that preceded the onset of the US-
UR and modeled the amplitude time source of the behavioral CR, just as did the
neuronal response in the hippocampus (McCormick et al. 1983). The following
brain areas showed this neuronal response predictive of the learned behavioral
CR: the relevant motor nuclei (as noted above), a region in the vicinity of the fifth
nucleus, various reticular regions, the cerebellar cortex (ansiform and anterior
lobes), the cerebellar interpositus nucleus, the pontine nuclei, and the red nucleus.
The response in the interpositus nucleus was particularly robust (Figure 4).
The fact that a neuronal model of the behavioral CR developed in the cerebellum
was suggestive of a memory trace but we knew from our earlier work on the hip-
pocampus that neuronal recordings, per se, could not identify the essential memory
trace. We completed a series of lesion studies, initially involving large cerebellar
lesions and later lesions limited to the interpositus nucleus (e.g., McCormick &
Thompson 1984a,b; Clark et al. 1984), all of which completely prevented learning
and completely and permanently abolished the CR with no effect on the UR (thus
ruling out performance variables).
In a long series of studies completed while we were at Stanford, we identi-
fied virtually the entire essential memory trace circuit using electrophysiological
recordings, electrical stimulation, lesions (aspiration, electrolytic, and chemical),
and anatomical pathway tracing methods (see Chapman et al. 1988; Christian &
Thompson 2003; McCormick et al. 1982a, 1985; Lavond et al. 1985; Mauk et al.
1986; Steinmetz et al. 1986, 1987; J.K. Thompson et al. 1985; Woodruff-Pak et al.
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12 THOMPSON
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Figure 4 Neuronal model of learned eyeblink response recorded from the cerebellar in-
terpositus nucleus. Each graph shows nictitating membrane movement (top trace) and a
histogram of multiple unit activity (bottom trace) averaged and summed over 100 trials.
Animals that received explicitly unpaired presentations of the conditioning stimuli do not
develop altered patterns of neuronal activity (left column). Trained animals develop a neu-
ronal model of the learned behavior. Total trace duration 750 msec. (From McCormick &
Thompson 1984b.)

1985). In brief, the efferent CR pathway projects from the interpositus nucleus ip-
silateral to the trained eye, via the superior cerebellar peduncle, to the contralateral
magnocellular red nucleus and to the relevant motor nuclei ipsilateral to the trained
eye. The cerebellar interpositus lesion abolition of the eyeblink CR is strictly ip-
silateral and is due to damage to neuron somas, not fibers of passage. The inferior
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LOCALIZATION OF MEMORY TRACES 13

olive-climbing fiber system appears to be the critical US reinforcing or teaching

pathway and the pontine nuclei and mossy fiber system appears to convey the
necessary CS information to the cerebellum.
A particularly satisfying aspect of our discovery of the essential role of the cere-
bellum in classical conditioning of discrete responses is the relevance of our work
to the human condition. Irene Daum and associates, working in Germany, repli-
cated in humans the fact that appropriate cerebellar damage completely prevents
learning of the eyeblink CR (Daum et al. 1993), since replicated in many other
studies. Christine Logan (a former graduate student of mine) and Scott Grafton, a
neurologist then at the University of Southern California, completed an extensive
positron emission tomography (PET) study of eyeblink conditioning in humans.
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They found significant activation in the cerebellar interpositus nucleus and several
loci in cerebellar cortex, in close agreement with our recording studies in the rabbit
cerebellum (Logan & Grafton 1995), again replicated in many other studies.
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A point that is obvious from our studies but seems not to be widely understood
is that our results on the role of the cerebellum in classical conditioning apply to the
learning of any discrete movement: eyeblink, head turn, forelimb flexion, hind-
limb flexion, etc. (see Shinkman et al. 1996). Eyeblink conditioning is simply
a convenient response to measure. Our findings to date seem to have identified
perhaps the critical function of the cerebellum, namely the learning of discrete
skilled movements, a basic notion proposed in classic theories of the cerebellum
as a learning machine (see Albus 1971, Eccles 1977, Ito 1984, Marr 1969). Indeed,
our work constitutes a compelling verification of these theories.
Our data to this point strongly supported the hypothesis that the essential mem-
ory trace was formed and stored in the cerebellum, but did not prove it. Hence we
adapted use of methods of reversible inactivation to localize the memory trace.
The logic is straightforward. Having identified the essential memory trace circuit,
reversibly inactivate each key locus in the circuit during training. If this completely
prevents learning at a given locus, then this locus either conveys essential affer-
ent information to the memory trace or is the site of the memory trace. However,
if reversible inactivation of a given locus does not prevent learning at all, then
this locus is efferent from the memory trace. But remember that inactivation of
all these essential loci will completely prevent expression of the CR. We infused
muscimol for inactivation of neuron cell bodies—it acts on gamma amino butyric
acid (GABAA) receptors as an agonist and completely shuts down (hyperpolarizes)
the neurons for several hours, after which they fully recover. To inactivate axons
we infused tetrodotoxin (TTX) (see, e.g., Krupa et al. 1993; Krupa & Thompson
1995, 1997; Krupa et al. 1996; Thompson & Krupa 1994). David Lavond and his
students completed parallel studies using reversible cooling (e.g., Clark et al. 1992,
Clark & Lavond 1993).
Results are completely consistent and have since been replicated in other labora-
tories. In brief (see Figure 5) inactivation limited to the anterior interpositus nucleus
completely prevented learning. After removal of inactivation animals showed no
signs of learning; with subsequent training they learned normally as though they
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14 THOMPSON
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Figure 5 Highly simplified schematic of the essential memory trace circuit for delay clas-
sical conditioning of the eyeblink response to illustrate the use of reversible inactivation to
localize memory traces. Shading in a, b, and c indicate reversible inactivation of the key re-
gion of each structure during training using muscimol; d indicates inactivation of the axonal
pathway exiting from the cerebellum, the superior cerebellar peduncle, using tetrodotoxin.
Inactivation of the interpositus (c) completely prevents learning, but inactivation of the su-
perior peduncle (d), the red nucleus (b), and the motor nuclei (a) does not prevent learning
at all. (Modified from Thompson & Krupa 1994.)

had no prior training; there was no savings. In complete contrast, inactivation of the
superior peduncle (the immediate efferent projection from the interpositus/dentate
nuclei), the red nucleus, and the relevant motor nuclei, although completely pre-
venting expression of the CR, did not prevent learning at all. After removal of
the inactivation the animals had fully learned to asymptote. Thus, inactivation
localized to the anterior interpositus nucleus completely prevented learning but in-
activation of its immediate output pathway did not prevent learning at all. The fact
that after interpositus inactivation there was no savings argues strongly that no part
of the memory trace developed in structures afferent to the interpositus. Similarly,
the fact that inactivation of structures efferent from the interpositus did not prevent
learning at all argues that no part of the memory is formed in these structures.
I noted above that several loci in the brain exhibited the neuronal model of the
learned behavioral CR, e.g., trigeminal nuclear region, pontine nuclei, etc. In a
series of studies Lavond and associates (and we) showed that with inactivation of
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LOCALIZATION OF MEMORY TRACES 15

the interpositus nucleus, all these models disappeared, i.e., they are all driven from
the interpositus (Bao et al. 2000, Lavond & Cartford 2000).
Because muscimol acts only on neuron cell bodies and not on axons, the in-
activation of the interpositus nucleus does not prevent normal activation of cere-
bellar cortex by the two major afferent systems, climbing fibers from the inferior
olive and mossy fibers from many sources. The fact that no learning at all oc-
curred with inactivation limited to the interpositus nucleus would seem to argue
against formation of memory traces in the cerebellar cortex. However, this inacti-
vation blocks the direct projections from the interpositus nucleus to the cerebellar
cortex.
When we first discovered the essential role of the cerebellum in eyeblink condi-
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tioning I had hoped the memory traces would be formed and stored in the cerebellar
cortex. The cellular machinery in the cortex is vast (each Purkinje neuron receives
up to 200,000 synapses from granule neurons). However, we were never able to
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completely prevent or abolish the CR by lesions limited to the cerebellar cortex.

Other workers, particularly Yeo and associates (Yeo et al. 1984), claimed to have
done so. The problem is with the lesion method—it is impossible to remove all
of the cerebellar cortex without damaging the critical region of the interpositus
nucleus (see detailed discussion in Christian & Thompson 2003). However, we
consistently found that with large cerebellar cortical lesions or reversible inactiva-
tion, learning was much slower and to a lesser degree and adaptive timing of the
CR was lost. Normally the eyeblink closure CR peaks at the onset of the US; it is
a maximally adaptive response. After large cortical lesions, the CR peaks earlier
in time and is no longer adaptive (Garcia et al. 1999, McCormick & Thompson
1984b). So the cortex is critically important for normal adaptive learning and it
seemed very likely that higher-order memory traces were established there. But
again the lesion method is inconclusive.
Fortunately nature provided us with an ideal preparation, the Purkinje cell
degeneration (pcd) mutant mouse. The brain of this mutant develops normally
until about two weeks after birth. Then over the next several weeks, all Purkinje
neurons in the cerebellar cortex die; as a result the animals have no functional
cerebellar cortex. But the interpositus nucleus remains functional. Results were
clear: The pcd mice were able to learn (L. Chen et al. 1996). They learned much
more slowly and to a lesser degree than normal mice of the same strain (littermates),
but they still showed significant and substantial learning. The CRs were shorter
latency in the pcd mice. They also showed rapid extinction with CS-alone training.
The possibility that the memory trace was somehow formed in loci other than the
cerebellum was ruled out by lesioning the interpositus nucleus bilaterally in pcd
mice before training. The lesioned pcd mice were unable to learn the conditioned
eyeblink response (L. Chen et al. 1999).
These results were very similar to the effects of large cerebellar cortical lesions
we had found earlier in rabbits. So the cortex is critically important for normal
learning, and higher-order memory traces are likely formed there. There is con-
siderable evidence supporting the view that a process of long-term potentiation
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16 THOMPSON

(LTD) at the parallel fiber-Purkinje neuron dendritic synapses, discovered by Ito

(see 1984), plays a key role in initial plasticity (C. Chen & Thompson 1995, C.
Chen et al. 1995, Shibuki et al. 1996, Kim & Thompson 1997). Indeed, Purk-
inje neurons show substantial changes in CS-evoked discharge frequency over the
course of training, with many showing learning-induced decrease in discharge fre-
quency, as could be expected if LTD developed (Christian et al. 2002, Christian
& Thompson 2003). Other lines of evidence also support the view that memory
traces are formed in cerebellar cortex (Cooke et al. 2004).
The evidence is now very strong that the basic essential memory trace is formed
and stored in the anterior interpositus nucleus for classical conditioning of the eye-
blink response. Indeed, the reversible inactivation studies provide the key evidence
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(Figure 5). It is clear that long-lasting changes in the critical neurons in the anterior
interpositus nucleus do develop. Thus, infusion of protein synthesis inhibitors in
this locus completely prevents learning (Bracha et al. 1998, G. Chen & Steinmetz
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2000, Gomi et al. 1999). Further, training results in selective expression of a protein
kinase in this locus, KKIAMRE, an enzyme for the cell division cycle (Gomi et al.
1999). Pharmacological isolation of the interpositus from the cerebellar cortex re-
veals clear learning-induced increases in excitability of interpositus neurons (Bao
et al. 2002, Garcia & Mauk 1998). Perhaps most convincing, eyeblink conditioning
results in a dramatic and highly significant increase in the number of excitatory
synapses (but not inhibitory synapses) in the interpositus nucleus (Kleim et al.
2002).

CONCLUSION
The essential circuitry for classical conditioning of the eyeblink response is shown
in Figure 6, along with the site of memory trace formation in the interpositus
nucleus and a putative site of plasticity in the cerebellar cortical neurons. The nature
of the memory is defined by this circuit; the circuit is the memory. The CS activates
the sensory afferent pathways to the site(s) of trace storage in the cerebellum,
which activates the efferent pathways to the motor nuclei and the learned behavior.
The content of the memory, the conditioned eyeblink response, is completely
defined and completely predictable from the essential circuit.
The formal criteria developed by Richard Morris and associates to demonstrate
that a given set of phenomena establish what they term the “synaptic plasticity and
memory” (SPM) hypothesis are as follows (Martin et al. 2000):

1. DETECTABILITY: If an animal displays memory of some previous expe-

rience, a change in synaptic efficacy should be detectable somewhere in its
nervous system.
2. MIMICRY: Conversely, if it were possible to induce the same spatial pattern
of synaptic weight changes artificially, the animal should display “apparent”
memory for some past experience which did not in practice occur.
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LOCALIZATION OF MEMORY TRACES 17

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Figure 6 Simplified schematic (most interneurons omitted) of the putative essential circuitry
for delay classical conditioning of eyeblink (and other discrete responses) learned with an
aversive US. (The sensory and motor nuclei activated depend of course on the nature of the
CS and US; the more central portions of the circuit appear to be general.) The reflex US-UR
pathway involves direct and indirect projection from the trigeminal nucleus to the motor nuclei
(for the eyeblink UR and CR, primarily accessory 6 and 7). The tone CS pathway projects
from auditory nuclei to the pontine nuclei and to the cerebellum as mossy fibers. The US
pathway includes projections from the trigeminal to the inferior olive and to the cerebellum
as climbing fibers. The CR pathway projects from the interpositus to the red nucleus and on
to premotor and motor nuclei. There is also a direct GABAergic inhibitory projection from
the interpositus to the inferior olive. Solid cell bodies and bar terminals indicate inhibitory
neurons; open cell bodies and fork terminals indicate excitatory neurons. Stars indicate sites
of plasticity based on current evidence. See text for details. (From Christian & Thompson
2003.) CR, conditioned response; CS, conditioned stimulus; UR, unconditioned response;
US, unconditioned stimulus.

3. ANTEROGRADE ALTERATION: Interventions that prevent the induction

of synaptic weight changes during a learning experience should impair the
animal’s memory of that experience.
4. RETROGRADE ALTERATION: Interventions that alter the spatial distri-
bution of synaptic weights induced by a prior learning experience (see de-
tectability) should alter the animal’s memory of that experience (p. 651).
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18 THOMPSON

I would expand the SPM notion to include nonsynaptic mechanisms of plasticity

and memory as well, referring to all as the memory trace. I would also note that
these criteria cannot be met unless the putative memory trace has been localized.
Evidence to date on localization of the memory trace(s) for classical condition-
ing of eyeblink and other discrete responses would seem to satisfy Morris’ criteria.
1. DETECTABILITY: There is a dramatic increase in neuronal/synaptic effi-
cacy in the cerebellum, both in the cortex of lobule HVI and in the anterior
interpositus nucleus as a result of training (see, e.g., Shinkman et al. 1996,
Bao et al. 2002). Indeed, long-term potentiation (LTP) has been reported to
occur in the interpositus nucleus (Racine et al. 1986).
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2. MIMICRY: Electrical microstimulation of the locus of the memory trace in

the anterior interpositus nucleus can evoke the response to be learned before
training (McCormick & Thompson 1984a, Chapman et al. 1988).
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3. ANTEROGRADE ALTERATION: Infusion of muscimol into the site of

the memory trace in the anterior interpositus nucleus completely prevents
learning, i.e., induction of the memory trace.
4. RETROGRADE ALTERATION: Infusion of muscimol into the site of the
memory trace in the anterior interpositus nucleus in a well-trained animal
alters the synaptic weights (shuts them down) and abolishes the animal’s
memory for that experience.
These of course are only opinions of what constitutes demonstration of a mem-
ory trace. In my view the key is that the essential circuit defines the memory, as I
noted above. But we still do not know the detailed nature of the memory trace in the
interpositus and how it is formed. It will be necessary to identify all the steps in the
causal chain from initial activation of the neurons at the beginning of training to the
final form of the memory trace, from the biochemical/genetic processes to the struc-
tural changes in the synapses and neurons that code the permanent memory trace.

ACKNOWLEDGMENTS
Work described in this paper was supported in part by National Science Foundation
Grant IBN 92,15069, National Institutes of Aging Grant AG14751, a grant from
the Sankyo Company, and funds from the University of Southern California.

The Annual Review of Psychology is online at http://psych.annualreviews.org

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Eyeblink Classical Conditioning: Animal Gould E, Beylin A, Tanapat P, Reeves A, Shors
Models, ed. DS Woodruff-Pak, JE Steinmetz, TJ. 1999. Learning enhances adult neurogen-
2:313–34. Boston, MA: Kluwer Acad. esis in the hippocampal formation. Nat. Neu-
Disterhoft JF, Olds J. 1972. Differential devel- rosci. 2:260–65
opment of conditioned unit changes in tha- Groves PM, Thompson RF. 1970. Habituation:
lamus and cortex of rat. J. Neurophysiol. a dual-process theory. Psychol. Rev. 77:419–
35:665–79 50
Eccles JC. 1977. An instruction-selection the- Hiraoka M, Shimamura M. 1977. Neural
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LOCALIZATION OF MEMORY TRACES 21

mechanisms of the corneal blinking reflex in Lavond DG, Cartford MC. 2000. Eyeblink
cats. Brain Res. 125:265–75 conditioning circuitry: tracing, lesion, and
Ito M. 1984. The Cerebellum and Neural Con- reversible lesion experiments. In Eyeblink
trol. New York: Raven Classical Conditioning: Animal Models, ed.
Ivkovich D, Thompson RF. 1997. Motor cortex DS Woodruff-Pak, JE Steinmetz, 2:51–80.
lesions do not affect learning or performance Boston, MA: Kluwer Acad.
of the eyeblink response in rabbits. Behav. Lavond DG, Hembree TL, Thompson RF. 1985.
Neurosci. 111:727–38 Effect of kainic acid lesions of the cerebellar
Kandel ER. 1975. The Cellular Basis of Behav- interpositus nucleus on eyelid conditioning
ior: An Introduction to Behavioral Neurobi- in the rabbit. Brain Res. 326:179–83
ology. San Francisco: Freeman Lavond DG, Kim JJ, Thompson RF. 1993.
Kandel ER, Spencer WA. 1968. Cellular neu- Mammalian brain substrates of aversive
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rophysiological approaches in the study of classical conditioning. Annu. Rev. Psychol.

learning. Physiol. Rev. 48:65–134 44:317–42
Kim JJ, Clark RE, Thompson RF. 1995. Lavond DG, Logan CG, Sohn JH, Garner WD,
by Ball State University on 01/05/09. For personal use only.

Hippocampectomy impairs the memory Kanzawa SA. 1990. Lesions of the cerebellar
of recently, but not remotely, acquired interpositus nucleus abolish both nictitating
trace eyeblink conditioned responses. Behav. membrane and eyelid EMG conditioned re-
Neurosci. 109:195–203 sponses. Brain Res. 514:238–48
Kim JJ, Thompson RF. 1997. Cerebellar cir- Logan CG, Grafton ST. 1995. Functional
cuits and synaptic mechanisms involved in anatomy of human eyeblink conditioning
classical eyeblink conditioning. Trends Neu- determined with regional cerebral glucose
rosci. 20:177–81 metabolism and positron-emission tomogra-
Kleim JA, Freeman JH Jr, Bruneau R, Nolan phy. Proc. Natl. Acad. Sci. USA 92:7500–4
BC, Cooper NR, Zook A, et al. 2002. Synapse Marr D. 1969. A theory of cerebellar cortex. J.
formation is associated with memory storage Physiol. 202:437–70
in the cerebellum. Proc. Natl. Acad. Sci. USA Martin SJ, Grimwood PD, Morris RGM. 2000.
99:13228–31 Synaptic plasticity and memory: an evalua-
Krupa DJ, Thompson JK, Thompson RF. 1993. tion of the hypothesis. Annu. Rev. Neurosci.
Localization of a memory trace in the mam- 23:649–711
malian brain. Science 260:989–91 Mauk MD, Steinmetz JE, Thompson RF. 1986.
Krupa DJ, Thompson RF. 1995. Inactivation of Classical conditioning using stimulation of
the superior cerebellar peduncle blocks ex- the inferior olive as the unconditioned stim-
pression but not acquisition of the rabbit’s ulus. Proc. Natl. Acad. Sci. USA 83:5349–53
classically conditioned eyeblink response. McCormick DA, Clark GA, Lavond DG,
Proc. Natl. Acad. Sci. USA 92:5097–101 Thompson RF. 1982a. Initial localization of
Krupa DJ, Thompson RF. 1997. Reversible in- the memory trace for a basic form of learning.
activation of the cerebellar interpositus nu- Proc. Natl. Acad. Sci. USA 79:2731–42
cleus completely prevents acquisition of the McCormick DA, Lavond DG, Thompson RF.
classically conditioned eyeblink response. 1982b. Concomitant classical conditioning
Learn. Mem. 3:545–56 of the rabbit nictitating membrane and eye-
Krupa DJ, Weng J, Thompson RF. 1996. In- lid responses: correlations and implications.
activation of brainstem motor nuclei blocks Physiol. Behav. 28:769–75
expression but not acquisition of the rabbit’s McCormick DA, Lavond DG, Thompson RF.
classically conditioned eyeblink response. 1983. Neuronal responses of the rabbit
Behav. Neurosci. 110:219–27 brainstem during performance of the clas-
Lashley KS. 1950. In search of the engram. Soc. sically conditioned nictitating membrane
Exp. Biol. Symp. 4:454–82 (NM/eyelid response). Brain Res. 271:73–88
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22 THOMPSON

McCormick DA, Steinmetz JE, Thompson RF. (Oryctolagus cuniculus). J. Comp. Physiol.
1985. Lesions of the inferior olivary com- Psychol. 79:328–33
plex cause extinction of the classically con- Shibuki K, Gomi H, Chen L, Bao S, Kim JJ,
ditioned eyeblink response. Brain Res. 359: et al. 1996. Deficient cerebellar long-term de-
120–30 pression, impaired eyeblink conditioning and
McCormick DA, Thompson RF. 1984a. Cere- normal motor coordination in GFAP mutant
bellum: essential involvement in the classi- mice. Neuron 16:587–99
cally conditioned eyelid response. Science Shinkman PG, Swain RA, Thompson RF. 1996.
223:296–99 Classical conditioning with electrical stim-
McCormick DA, Thompson RF. 1984b. Neu- ulation of cerebellum as both conditioned
ronal responses of the rabbit cerebellum dur- and unconditioned stimulus. Behav. Neu-
ing acquisition and performance of a clas- rosci. 110:914–21
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sically conditioned nictitating membrane- Shors TJ, Matzel LD. 1997. Long-term po-
eyelid response. J. Neurosci. 4:2811–22 tentiation: What’s learning got to do with
McGlinchey-Berroth R, Carrillo MC, Gabrieli it? Behav. Brain Sci. 20:597–614; discussion
by Ball State University on 01/05/09. For personal use only.

JD, Brawn CM, Disterhoft JF. 1997. Im- 614–55

paired trace eyeblink conditioning in bilat- Shors TJ, Miesegaes G, Beylin A, Zhao M,
eral, medial-temporal lobe amnesia. Behav. Rydel T, Gould E. 2001. Neurogenesis in the
Neurosci. 111:873–82 adult is involved in the formation of trace
Moyer JR Jr, Deyo RA, Disterhoft JF. 1990. memories. Nature 410:372–76
Hippocampectomy disrupts trace eye-blink Solomon PR, Vander Schaaf ER, Thompson
conditioning in rabbits. Behav. Neurosci. RF, Weisz DJ. 1986. Hippocampus and trace
104:243–52 conditioning of the rabbit’s classically condi-
Olds J, Anderson ML, McPhie DL, Staten tioned nictitating membrane response. Behav
LD, Alkon DL. 1989. Imaging of memory- Neurosci. 100:729–44
specific changes in the distribution of pro- Squire LR. 1987. Memory and Brain. New
tein kinase C in the hippocampus. Science York: Oxford Univ. Press
245:866–69 Squire LR. 1992. Declarative and non-
Olds J, Disterhoft J, Segal M, Kornblith DL, declarative memory: multiple brain systems
Hirsh R. 1972. Learning centers of rat brain supporting learning and memory. J. Cogn.
mapped by measuring latencies of condi- Neurosci. 4:232–43
tioned unit responses. J. Neurophysiol. 35: Steinmetz JE, Logan CG, Rosen DJ, Thompson
202–19 JK, Lavond DG, Thompson RF. 1987. Ini-
Patterson MM, Cegavske CF, Thompson RF. tial localization of the acoustic conditioned
1973. Effects of classical conditioning stimulus projection system to the cerebel-
paradigm on hindlimb flexor nerve response lum essential for classical eyelid condition-
in immobilized spinal cat. J. Comp. Physiol. ing. Proc. Natl. Acad. Sci. USA 84:3531–35
Psychol. 84:88–97 Steinmetz JE, Rosen DJ, Chapman PF, Lavond
Racine RJ, Wilson DA, Gingell R, Sunderland DG, Thompson RF. 1986. Classical condi-
D. 1986. Long-term potentiation in the inter- tioning of the rabbit eyelid response with a
positus and vestibular nuclei in the rat. Exp. mossy fiber stimulation CS. I. Pontine nuclei
Brain Res. 63:158–62 and middle cerebellar peduncle stimulation.
Schacter DL. 1987. Implicit memory: history Behav. Neurosci. 100:871–80
and current status. Exp. Psychol. Learn. Thompson JK, Lavond DG, Thompson RF.
Mem. Cogn. 13:501–18 1985. Cerebellar interpositus/dentate nuclei
Schmaltz LW, Theios J. 1972. Acquisition afferent seen with retrograde fluorescent trac-
and extinction of a classically conditioned ers in the rabbit. Neurosci. Abstr. 11:1112
response in hippocampectomized rabbits Thompson RF. 1997. Classical conditioning
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LOCALIZATION OF MEMORY TRACES 23

has much to do with LTP. Behav. Brain Sci. sponse: I. Primary Field (AI). Behav. Neu-
20:632–33 rosci. 98:171–88
Thompson RF, Berger TW, Cegavske CF, Pat- Weisz DJ, Clark GA, Thompson RF. 1984.
terson MM, Roemer RA, et al. 1976. The Increased activity of dentate granule cells
search for the engram. Am. Psychol. 31:209– during nictitating membrane response condi-
27 tioning in rabbits. Behav. Brain Res. 12:145–
Thompson RF, Kim JJ. 1996. Memory systems 54
in the brain and localization of a memory. Wilson MA, Tonegawa S. 1997. Synaptic plas-
Proc. Natl. Acad. Sci. USA 93:13438–44 ticity, place cells and spatial memory: study
Thompson RF, Krupa DJ. 1994. Organization with second generation knockouts. Trends
of memory traces in the mammalian brain. Neurosci. 20:102–6
Annu. Rev. Neurosci. 17:519–49 Woodruff-Pak DS, Lavond DG, Thompson RF.
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Thompson RF, Patterson MM, Teyler TJ. 1972. 1985. Trace conditioning: abolished by cere-
Neurophysiology of learning. Annu. Rev. bellar nuclear lesions but not lateral cerebel-
Psychol. 23:73–104 lar cortex aspirations. Brain Res. 348:249–
by Ball State University on 01/05/09. For personal use only.

Thompson RF, Spencer WA. 1966. Habituation: 60

a model phenomenon for the study of neu- Woody CD, Alkon DL, Hay B. 1984. Depolar-
ronal substrates of behavior. Psychol. Rev. ization-induced effects of Ca2+-calmodulin-
73:16–43 dependent protein kinase injection, in vivo,
Tulving E. 1985. How many memory systems in single neurons of cat motor cortex. Brain
are there? Am. Psychol. 40:385–98 Res. 321:192–97
Wagner AR, Donegan NH. 1989. Some re- Yeo CH, Hardiman MJ, Glickstein M. 1984.
lationships between a computational model Discrete lesions of the cerebellar cortex abol-
(SOP) and a neural circuit for Pavlovian ish the classically conditioned nictitating
(rabbit eyeblink) conditioning. In The Psy- membrane response of the rabbit. Behav.
chology of Learning and Motivation, ed. Brain Res. 13:261–66
RD Hawkins, GH Bower, 22:157–203. San Young RA, Cegavske CF, Thompson RF.
Diego: Academic 1976. Tone-induced charges in excitabil-
Weinberger NM, Hopkins W, Diamond DM. ity of abducens motoneurons and the reflex
1984. Physiological plasticity of single neu- path of the rabbit nictitating membrane re-
rons in auditory cortex of the cat during sponse. J. Comp. Physiol. Psychol. 90:424–
acquisition of the papillary conditioned re- 34
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December 8, 2004 12:13 Annual Reviews AR231-FM

Annual Review of Psychology

Volume 56, 2005

CONTENTS
Frontispiece—Richard F. Thompson xviii
PREFATORY
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In Search of Memory Traces, Richard F. Thompson 1

DECISION MAKING
Indeterminacy in Brain and Behavior, Paul W. Glimcher 25
by Ball State University on 01/05/09. For personal use only.

BRAIN IMAGING/COGNITIVE NEUROSCIENCE

Models of Brain Function in Neuroimaging, Karl J. Friston 57
MUSIC PERCEPTION
Brain Organization for Music Processing, Isabelle Peretz
and Robert J. Zatorre 89
SOMESTHETIC AND VESTIBULAR SENSES
Vestibular, Proprioceptive, and Haptic Contributions
to Spatial Orientation, James R. Lackner and Paul DiZio 115
CONCEPTS AND CATEGORIES
Human Category Learning, F. Gregory Ashby and W. Todd Maddox 149
ANIMAL LEARNING AND BEHAVIOR: CLASSICAL
Pavlovian Conditioning: A Functional Perspective,
Michael Domjan 179
NEUROSCIENCE OF LEARNING
The Neuroscience of Mammalian Associative Learning,
Michael S. Fanselow and Andrew M. Poulos 207
HUMAN DEVELOPMENT: EMOTIONAL, SOCIAL, AND PERSONALITY
Behavioral Inhibition: Linking Biology and Behavior Within a
Developmental Framework, Nathan A. Fox, Heather A. Henderson,
Peter J. Marshall, Kate E. Nichols, and Melissa A. Ghera 235
BIOLOGICAL AND GENETIC PROCESSES IN DEVELOPMENT
Human Development: Biological and Genetic Processes,
Irving I. Gottesman and Daniel R. Hanson 263

vii
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December 8, 2004 12:13 Annual Reviews AR231-FM

viii CONTENTS

SPECIAL TOPICS IN PSYCHOPATHOLOGY

The Psychology and Neurobiology of Suicidal Behavior,
Thomas E. Joiner Jr., Jessica S. Brown, and LaRicka R. Wingate 287
DISORDERS OF CHILDHOOD
Autism in Infancy and Early Childhood, Fred Volkmar,
Kasia Chawarska, and Ami Klin 315
CHILD/FAMILY THERAPY
Youth Psychotherapy Outcome Research: A Review and Critique
of the Evidence Base, John R. Weisz, Amanda Jensen Doss,
Annu. Rev. Psychol. 2005.56:1-23. Downloaded from arjournals.annualreviews.org

and Kristin M. Hawley 337

ALTRUISM AND AGGRESSION
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Prosocial Behavior: Multilevel Perspectives, Louis A. Penner,

John F. Dovidio, Jane A. Piliavin, and David A. Schroeder 365
INTERGROUP RELATIONS, STIGMA, STEREOTYPING,
PREJUDICE, DISCRIMINATION
The Social Psychology of Stigma, Brenda Major
and Laurie T. O’Brien 393
PERSONALITY PROCESSES
Personality Architecture: Within-Person Structures and Processes,
Daniel Cervone 423
PERSONALITY DEVELOPMENT: STABILITY AND CHANGE
Personality Development: Stability and Change, Avshalom Caspi,
Brent W. Roberts, and Rebecca L. Shiner 453
WORK MOTIVATION
Work Motivation Theory and Research at the Dawn of the Twenty-First
Century, Gary P. Latham and Craig C. Pinder 485
GROUPS AND TEAMS
Teams in Organizations: From Input-Process-Output Models to IMOI
Models, Daniel R. Ilgen, John R. Hollenbeck, Michael Johnson,
and Dustin Jundt 517
LEADERSHIP
Presidential Leadership, George R. Goethals 545
PERSONNEL EVALUATION AND COMPENSATION
Personnel Psychology: Performance Evaluation and Pay for Performance,
Sara L. Rynes, Barry Gerhart, and Laura Parks 571
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December 8, 2004 12:13 Annual Reviews AR231-FM

CONTENTS ix

PSYCHOPHYSIOLOGICAL DISORDERS AND PSYCHOLOGICAL EFFECTS

ON MEDICAL DISORDERS
Psychological Approaches to Understanding and Treating Disease-Related
Pain, Francis J. Keefe, Amy P. Abernethy, and Lisa C. Campbell 601
TIMELY TOPIC
Psychological Evidence at the Dawn of the Law’s Scientific Age,
David L. Faigman and John Monahan 631

INDEXES
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Subject Index 661

Cumulative Index of Contributing Authors, Volumes 46–56 695
Cumulative Index of Chapter Titles, Volumes 46–56 700
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ERRATA
An online log of corrections to Annual Review of Psychology chapters
may be found at http://psych.annualreviews.org/errata.shtml
 18 Nov 2004 10:44 AR AR231-PS56-02.tex AR231-PS56-02.sgm LaTeX2e(2002/01/18) P1: IKH
10.1146/annurev.psych.55.090902.141429

Annu. Rev. Psychol. 2005. 56:25–56

doi: 10.1146/annurev.psych.55.090902.141429
Copyright
c 2005 by Annual Reviews. All rights reserved
First published online as a Review in Advance on September 10, 2004

INDETERMINACY IN BRAIN AND BEHAVIOR

Paul W. Glimcher
Center for Neural Science, New York University, New York, New York 10003;
email: [email protected]
Annu. Rev. Psychol. 2005.56:25-56. Downloaded from arjournals.annualreviews.org

Key Words randomness, decision-making, choice, game theory

■ Abstract The central goal of modern science that evolved during the Enlight-
enment was the empirical reduction of uncertainty by experimental inquiry. Although
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there have been challenges to this view in the physical sciences, where profoundly
indeterminate events have been identified at the quantum level, the presumption that
physical phenomena are fundamentally determinate seems to have defined modern be-
havioral science. Programs like those of the classical behaviorists, for example, were
explicitly anchored to a fully deterministic worldview, and this anchoring clearly in-
fluenced the experiments that those scientists chose to perform. Recent advances in the
psychological, social, and neural sciences, however, have caused a number of scholars
to begin to question the assumption that all of behavior can be regarded as fundamen-
tally deterministic in character. Although it is not yet clear whether the generative mech-
anisms for human and animal behavior will require a philosophically indeterminate
approach, it is clear that behavioral scientists of all kinds are beginning to engage the
issues of indeterminacy that plagued physics at the beginning of the twentieth century.

CONTENTS
INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Determinism and the Philosophy of Science . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Do Indeterminacies in the Physical World Matter for Behavioral
Scientists? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
THE RISING TIDE OF APPARENT INDETERMINACY . . . . . . . . . . . . . . . . . . . . . 29
Indeterminacy in the Social Sciences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Empirical Measurements of Behavioral Indeterminacy . . . . . . . . . . . . . . . . . . . . . . 35
Reducing Uncertainty: Looking for Determinacy with Neurophysiology . . . . . . . . 40
Indeterminacy at the Cellular and Subcellular Levels . . . . . . . . . . . . . . . . . . . . . . . . 46
THE CHALLENGE OF INDETERMINACY FOR BEHAVIORAL
SCIENCE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

INTRODUCTION
Our modern view that the central function of scientific inquiry is to reduce uncer-
tainty emerged early in the scientific revolution that constituted the Enlightenment;
by the time of Galileo’s death (cf. Bacon 1620, Descartes 1637, Galilei 1630,
0066-4308/05/0203-0025$14.00 25
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26 GLIMCHER

Kepler 1618–1621) it was clear that improving the accuracy with which one could
predict future events as determinate processes would be a central feature of the
scientific method at both theoretical and empirical levels in the physical sciences.
Over the course of the eighteenth and nineteenth centuries, the early social sci-
ences emulated this trend, seeking to develop causal relationships in a testable and
determinate fashion (cf. Keynes 1936, Smith 1776). By the twentieth century, the
notion that scientific inquiry would reduce animal behavior to deterministic cer-
tainty had become a mainstream notion in psychological circles as well. Nowhere
is this clearer than in the work of Pavlov. As Pavlov put it in Conditioned Reflexes:

The physiologist must thus take his own path, where a trail has already been
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blazed for him. Three hundred years ago Descartes evolved the idea of the
reflex. Starting from the assumption that animals behaved simply as machines,
he regarded every activity of the organism as a necessary reaction to some
by Ball State University on 01/05/09. For personal use only.

external stimulus. . .. A bold attempt to apply the idea of the reflex to the
activities of the [cerebral] hemispheres was made by the Russian physiologist
I.M. Sechenov, on the basis of the knowledge available in his day of the
physiology of the central nervous system. In a pamphlet entitled “Reflexes of
the Brain,” published in Russia in 1863, he attempted to represent the activities
of the cerebral hemispheres as reflex—that is to say, as determined. (Pavlov
1927)

In the period that followed, Skinner and his students (cf. Skinner 1938) strength-
ened this notion, and psychologists as a group largely embraced the idea that a
complete psychological theory would be a determinate one. By studying the causal
relationships between environment, organism, and response, these scientists be-
gan the process of developing a predictive and testable theory of psychology.
The twentieth century witnessed a similar trend in the effective application of the
scientific method toward understanding the biological sources of behavior, and
as a result, saw the development of a powerful deterministic program for under-
standing biological systems. Charles Sherrington (1906), for example, applied this
programmatic approach to the physiological study of reflexes with great effect.

Determinism and the Philosophy of Science

In philosophical circles, the central role of a determinate worldview in the classical
scientific method also became a formalized principle in this period. In the early part
of the twentieth century, the philosopher Karl Popper (1934) explicitly defined the
goal of modern science as the falsification of extant theories through experimental
inquiry. For Popper, theories could never be proven in practice, only subjected to
the test of falsification.
If experimental evidence falsifies a theory then it can be discarded; if exper-
imental evidence corroborates a theory then it can be tentatively retained. What
is critical about this logic is what it implies about indeterminacy. Consider the
theoretical claim that if I flip a certain coin there is a 50% chance it will land
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INDETERMINATE BEHAVIOR 27

heads-up. As Popper points out, this is not only an unverifiable theoretical claim,
but also an untestable one; my assertion predicts all possible empirical outcomes
and is thus unfalsifiable. Even more importantly, my theoretical claim predicts as
an experimental result all possible finite series of coin flips that could ever be ob-
served. If the coin is equally likely to land heads-up and tails-up, then any specific
series of heads and tails is equally likely, whether that be six heads in a row or six
flips that alternate between heads and tails. No formal prediction of any particu-
lar outcome is ever possible and for this reason Popper argued that probabilistic
claims about indeterminate processes were irremediably problematic. Indeed, in
his early writings Popper even used this to argue that the notion of a fundamentally
indeterminate universe is at base a nonscientific proposition.
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In the 1920s and 1930s, however, the emerging discipline of quantum physics
raised an important challenge to this notion that had evolved during the Enlight-
enment and motivated much of Popper’s work. Based initially on the work of
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Heisenberg and his colleagues (Heisenberg 1930, 1952), strong evidence arose
suggesting that several phenomena that occur at the atomic and subatomic scales
are, in fact, fundamentally indeterminate and thus could be described only proba-
bilistically. This was a critical challenge to the existing philosophy of science as
expressed by Popper because that philosophy argued that a theory of physics built
upon probability theory was unfalsifiable, perhaps even unscientific. Nevertheless,
the empirical evidence gathered during that period seemed to indicate unambigu-
ously that at a small enough scale of analysis, events occur that are fundamentally
indeterminate. This indicated that the philosophy of science, rather than the reality
of our physical universe, might have to change.

Do Indeterminacies in the Physical World Matter

for Behavioral Scientists?
What, if any, are the implications of these issues for the study of behavior? Even
if there are fundamental indeterminacies in the physical world, should this matter
to behavioral scientists? Many scholars believe that the quantum physicist Edwin
Schrodinger provided an answer to that question in his book, What Is Life (1944),
in which he argued that for any organism to survive it must operate, in principle,
in a fully determinate environment. Indeterminacy, he believed, would be lethal
to living systems. Schrodinger’s own work (cf. 1951) had demonstrated that at
the atomic and subatomic scales, matter can be described only in probabilistic
terms, but it had also shown that large aggregates of these elementary particles
behaved in an effectively determinate manner. His argument was that living cells
were large enough objects that they would never interact with single atomic or
subatomic particles, but only with these larger determinate aggregates. In essence,
he argued that cells were large enough that quantum fluctuations in the properties
of individual atoms would have no effect on them. Indeed, he went on to argue
that biological cells are the size that they are specifically because quantum inde-
terminacy prevents them from surviving if they become any smaller. Biologists,
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28 GLIMCHER

psychologists, and social scientists, he assured us, need not be concerned with
fundamental indeterminacy in the universe:

If it were not so, if we were organisms so sensitive that a single atom, or even
a few atoms, could make a perceptible impression on our senses—Heavens,
what would life be like! To stress one point: an organism of that kind would
most certainly not be capable of developing the kind of orderly thought which,
after passing through a long sequence of earlier stages, ultimately results in
forming, among many other ideas, the idea of an atom. (Schrodinger 1951)

Recently, however, evidence has begun to arise in the social, psychological,

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and neurobiological domains that suggests that, at larger scales of analysis than
the one Schrodinger examined in What Is Life, living systems exhibit behavior
that is apparently indeterminate (cf. Hastie 2001, Schall 2004, Shafir & LeBoeuf
2002, Staddon & Cerutti 2003). At the largest scale of analysis, social scientists
by Ball State University on 01/05/09. For personal use only.

working in areas such as the theory of games have begun to argue that for behavior
to be efficient under some circumstances, it must be irreducibly uncertain from the
point of view of other organisms and therefore must be studied with the tools of
probability theory. In principle, this raises critical problems for game theory. For all
of the reasons Popper identified, when game theory makes probabilistic predictions
it does so in a manner that is nonfalsifiable. Of course, if Schrodinger was correct,
the apparent indeterminacy of game theory presents only a temporary impediment
to scientific inquiry. A reductionist approach to human behavior during strategic
games would ultimately reveal the mechanisms that give rise to this apparent
indeterminacy and thus should ultimately yield a falsifiable determinate theory of
human behavior. Although contemporary game theory thus faces indeterminacy,
empirical science can hope to resolve this apparent indeterminacy by reduction.
Interestingly, however, psychologists working at a lower level of reduction than
social scientists have also begun to find evidence of apparent indeterminacy in
the systems they study (cf. Staddon & Cerruti 2003). Recently, psychologists
have begun the analysis of apparently stochastic patterns of individual responses
and have been able to demonstrate classes of individual behavior that appear to
be as fully random as can be measured. Indeterminacy, in the hands of these
psychologists, seems to be an apparent feature of the behavior of single humans
and animals. At a yet deeper level of reduction, neurobiologists have also begun
to gather evidence for the existence of apparently indeterminate processes within
the architecture of the mammalian brain (cf. Schall 2004). The patterns of action
potentials generated by individual neurons, for example, appear highly stochastic
for reasons that are not yet well understood.
Growing evidence that apparently indeterminate processes operate at social,
psychological, and even neurobiological levels are bringing behavioral scientists
face-to-face with the same philosophical and scientific issues faced by Popper,
Heisenberg, Schrodinger, and others in the last century. Can such theories be
scientific, or is calling a neural signal or a behavior a random process only an
excuse for ignorance? It may be that behavioral scientists will choose to assert as an
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INDETERMINATE BEHAVIOR 29

axiom that all of the physical phenomena we study are fundamentally determinate
in order to avoid these issues, but on the other hand such an assertion may force
us to neglect a growing body of compelling evidence.

THE RISING TIDE OF APPARENT INDETERMINACY

Indeterminacy in the Social Sciences
Like scholars in the physical sciences, social scientists in the eighteenth and nine-
teenth centuries strongly emphasized a determinate scientific approach in their
Annu. Rev. Psychol. 2005.56:25-56. Downloaded from arjournals.annualreviews.org

study of human behavior. The classic economic theory of that period, for example,
rested on the foundation of a theory of determinate utility developed by Blaise
Pascal (1670, Arnauld & Nicole 1662) and Daniel Bernoulli (1738). This utility
theory argued that humans act predictably to maximize benefits and to minimize
by Ball State University on 01/05/09. For personal use only.

costs, and that the costs and benefits of any action can be reliably computed. Pascal
had developed this basic logic in the seventeenth century, arguing that the “ex-
pected value” of an action was equal to the product of any possible gain or loss
incurred by that action and the likelihood of the gain or loss. Bernoulli had ex-
tended this notion with the observation that humans appear at an empirical level to
be more averse to risk than Pascal’s formulation predicts. Bernoulli’s conclusion
was that humans made decisions based on the product of a subjective estimate
of cost or benefit and the likelihood of that gain or loss, rather than based on an
objective measure of gains or losses. Because of the precise form of his hypoth-
esis, Bernoulli was able to show that this notion could successfully account for
the empirically observed aversion of humans to risk. Thus, the critical idea that
utility theory advanced was that one could compute the relative desirabilities of all
possible actions to a chooser and, except in the presumably rare case where two
actions have identical subjective desirabilities, one could then predict the actions
of a chooser with determinate precision. Building on this foundation, Adam Smith
(1776) argued that all economic actors could be seen as effectively trading off costs
and benefits to maximize gain in a complex marketplace. The prices of goods, for
example, were presumed to be set by the determinate interactions of supply and
demand curves that represented the aggregate subjective desirabilities and costs
of goods to consumers and producers. It was thus a central thesis of eighteenth-
and nineteenth-century economic theory that the rational process by which de-
sirability was assessed could be accurately modeled and that these models made
deterministic predictions about human behavior.
Importantly, the incorporation of likelihoods into expected utility theory al-
lowed the approach to make determinate predictions even when the environment
in which human decision makers operated was unpredictable. Choosers were as-
sumed to consider risk when they determined the desirability of an action, and the
theory explicitly and convincingly predicted that no feature of this environmental
uncertainty would be presumed to propagate into the behavior of the choosers.
The only time that utility theory predicted indeterminacy in behavior was when
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30 GLIMCHER

two or more mutually exclusive actions had precisely equal subjective desirabili-
ties, and the importance of that particular situation seemed limited to the classical
economists.
In the first half of the twentieth century, the theory of games developed by John
VonNeumann, Oskar Morgenstern, and John Nash directly challenged this deter-
minate approach (Nash 1950a,b, 1951; VonNeumann & Morgenstern 1944). Game
theory represented a deviation from the classical tradition specifically because it
proposed that when a rational chooser faces an intelligent and self-interested oppo-
nent, rather than a passive economic environment, situations could easily arise in
which the subjective desirabilities of two or more actions are driven toward precise
equality. The theory went on to make surprising and fundamentally indeterminate
Annu. Rev. Psychol. 2005.56:25-56. Downloaded from arjournals.annualreviews.org

predictions about how rational humans would behave under many conditions that
could be well described by game theory.
To understand this theoretical insight, consider two opponents repeatedly play-
by Ball State University on 01/05/09. For personal use only.

ing the childhood game of rock-paper-scissors in which the loser pays the winner
$2 on a round won by playing paper and $1 on a round won by playing scissors or
rock. If the behavior of one’s opponent is unpredictable, any response can win, in
principle. Paper will beat a play of rock for $2, scissors will beat a play of paper
for $1, and rock will beat a play of scissors, again for $1. Classical utility theory
assumes that humans choose between actions by multiplicatively combining the
subjective value and likelihood of each outcome and then selecting the action with
the outcome that yields the highest expected utility. Assuming naively that one’s
opponent is equally likely to play rock, or paper, or scissors, the greater value of
winning with paper should lead all players to select paper deterministically on
each round. What VonNeumann recognized was that this assumption about the
behavior of one’s opponent simply could not be correct. A competitor who simply
selected scissors could reliably defeat any player who actually behaved in accord
with this strategy.
Game theory, as developed by VonNeumann & Morgenstern (1944), addresses
this limitation of classical utility theory by making the assumption that both players
are aware that they face an intelligent opponent who can anticipate their actions
and that both players will shape their behavior accordingly to minimize losses and
maximize gains. To accomplish this, players must take into account the potential
payoffs associated with each choice, as specified by classical utility theory, but
they must also consider how the actions of their opponent will influence those
payoffs. Consider again the situation in rock-paper-scissors. Winning with paper
yields twice as much money as winning with rock or scissors, but deterministically
playing paper leads to certain defeat. What VonNeumann & Morgenstern showed
was that under these conditions we can predict that a rational player will titrate risk
against gain and play paper two-thirds of the time, scissors one-sixth of the time,
and rock one-sixth of the time. Critically, however, he must avoid making his two-
thirds, one-sixth, one-sixth selections in a determinate fashion; for example, in a
repeated version of the game by playing paper, then scissors, then paper, then rock,
then paper, and then paper. Were his opponent to divine the determinate nature
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INDETERMINATE BEHAVIOR 31

of such a strategy (through observation, for example), then winning would again
become trivial for that opponent. He would only have to play scissors, then rock,
then scissors, then paper, then scissors, and then scissors to assure a consistent win.
The only way to avoid this trap is for a player to incorporate apparent indeterminacy
directly into his behavior. He must in essence flip a weighted coin on each round
to select between rock and paper and scissors. VonNeumann & Morgenstern were
well aware of the implications of this observation. It suggested that under some
conditions the study of economic choice would have to become a probabilistic
process. As they put it:
Consider now a participant in a social exchange economy. His problem has,
Annu. Rev. Psychol. 2005.56:25-56. Downloaded from arjournals.annualreviews.org

of course, many elements in common with a maximum problem. [A problem

in which a single economic actor seeks to maximize his gain by classically
deterministic processes.] But it also contains some, very essential, elements
by Ball State University on 01/05/09. For personal use only.

of an entirely different nature. He too tries to obtain an optimum result. But in

order to achieve this, he must enter into relations of exchange with others. If
two or more persons exchange goods with each other, then the results for each
one will depend in general not merely upon his own actions but on those of
the others as well. Thus each participant attempts to maximize a function (his
above-mentioned “result”) of which he does not control all of the variables.
This is certainly no maximization problem, but a peculiar and disconcerting
mixture of several conflicting maximum problems. Every participant is guided
by another principle and neither determines all of the variables which affect
his interest.
This kind of problem is nowhere dealt with in classical mathematics. . ..
We hope that the reader will be convinced by the above that they face here and
now a really conceptual—and not merely technical—difficulty. And it is this
problem which the theory of “games of strategy” is mainly devised to meet.
(VonNeumann & Morgenstern 1944)
VonNeumann & Morgenstern’s critical insight was that under conditions of
this type choosers might not be able to identify a single course of action that
is deterministically optimal. Instead, they may be forced to select a course of
action in as random a fashion as possible. It is this strategy of random selection,
known now as a mixed strategy, that distinguishes VonNeumann & Morgenstern’s
approach from more classical deterministic approaches to the study of behavior.
In sum, VonNeumann & Morgenstern argued that human behavior, under some
conditions, must appear indeterminate in order to be efficient. They made this
point elegantly when they described, in game theoretic form, a conflict between
Sherlock Holmes and his archenemy, Professor Moriarity:
Sherlock Holmes desires to proceed from London to Dover and hence to the
continent in order to escape from Professor Moriarity who pursues him. Hav-
ing boarded the train he observes, as the train pulls out, the appearance of Pro-
fessor Moriarity on the platform. Sherlock Holmes takes it for granted—and
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32 GLIMCHER

in this he is assumed to be fully justified—that his adversary, who has seen

him, might secure a special train and overtake him. Sherlock Holmes is faced
with the alternative of going to Dover or of leaving the train at Canterbury, the
only intermediate station. His adversary—whose intelligence is assumed to
be fully adequate to visualize these possibilities—has the same choice. Both
opponents must choose the place of their detrainment in ignorance of the
other’s corresponding decision. If, as a result of these measures, they should
find themselves, in fine, on the same platform, Sherlock Holmes may with
certainty expect to be killed by Moriarity. If Holmes reaches Dover unharmed
he can make good his escape.
What are the good strategies, particularly for Sherlock Holmes? [Set the
Annu. Rev. Psychol. 2005.56:25-56. Downloaded from arjournals.annualreviews.org

value] to Professor Moriarity [of] catching Sherlock Holmes [at], say 100.
[Alternatively, consider what happens if] Sherlock Holmes successfully es-
caped to Dover, while Moriarity stopped at Canterbury. This is Moriarity’s
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defeat as far as the present action is concerned, and should be described by

a big negative value of the matrix element [for Moriarity]—in the order of
magnitude but smaller than the positive value mentioned above—say, −50.
[Finally, consider what happens if] Sherlock Holmes escapes Moriarity at the
intermediate station, but fails to reach the Continent. This is best viewed as a
tie, and assigned the matrix element 0.
[From a mathematical analysis of these values one can conclude that] the
good strategies (e for Moriarity, n for Sherlock Holmes) [are]:
e = {3/5, 2/5}, n = {2/5, 3/5}
Thus Moriarity should go to Dover with a probability of 60% while Sher-
lock Holmes should stop at the intermediate station with a probability of
60%, the remaining 40% being left in each case for the other alternative.1
(VonNeumann & Morgenstern 1944, pp. 177–178)
Of course, this theoretical formulation raises critical questions about the scien-
tific nature of game theory. If game theory predicts that Holmes will get off the
train at Canterbury with a 60% probability, any action Holmes takes is compatible
with the theory. VonNeumann & Morgenstern recognized this but were adamant
that this was still the only rational strategy for Holmes to adopt. Holmes must, they
argued, be as indeterminate as possible in selecting a course of action. In essence,
he must appear to have flipped a weighted coin (weighted 60% for Canterbury and
40% for Dover) in order to maximize his chance of survival. This was true, Von-
Neumann & Morgenstern suggested, irrespective of whether the theory of games
preserved Popperian falsifiability.
By the early 1950s, John Nash (1950a,b; 1951) had seen an interesting addi-
tional level of structure in game theoretic problems that required a mixed strategy,

1
Our result for e, n yields that Sherlock Holmes is as good as 48% dead when his train pulls
out from Victoria Station.
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INDETERMINATE BEHAVIOR 33

or apparently indeterminate, solution. Building on the work of VonNeumann &

Morgenstern, he concluded that stable mixed strategies must in principle reflect an
equilibrium point at which the subjective desirabilities of the two or more actions
being mixed were precisely equivalent. He argued that it was only this equivalence
that could produce the indeterminate behavior that VonNeumann & Morgenstern
had predicted. Consider again the situation when a single player must, on repeated
rounds, select rock or paper or scissors. If any one of these is truly preferable
as a choice, then we can assume that the chooser will always select that option.
Mixed strategies should thus emerge, Nash reasoned, only when the two or more
actions that are being mixed have identical average desirabilities. Working from
VonNeumann & Morgenstern’s insights, Nash argued that these two equivalent
Annu. Rev. Psychol. 2005.56:25-56. Downloaded from arjournals.annualreviews.org

desirabilities emerge when the competitive interactions of the two players drive
them toward an equilibrium at which the two or more actions being mixed are of
equal desirability. What Nash argued was that mixed-strategy equilibriums emerge
by Ball State University on 01/05/09. For personal use only.

from dynamic interactions between the players, which yield equal average desir-
abilities, and thus totally indeterminate patterns of behavioral choice. Classical
utility theory had presumed that situations in which two or more actions have
precisely equal subjective desirabilities would be encountered only rarely. Nash’s
insight was that not only are they encountered, but the dynamic interactions that
occur during strategic games actively create these situations of equal subjective
desirability.
From the point of view of indeterminacy, the critical insight that VonNeumann,
Morgenstern, and Nash offered was that indeterminacy is a requisite feature of
efficient behavior in a competitive world. That insight means either that humans
and animals appear indeterminate to each other under some conditions or they
behave inefficiently.
In 1982, the evolutionary biologist John Maynard Smith also engaged indeter-
minacy during strategic games, but from an evolutionary perspective. He argued
that any species involved in an internal competition for resources could be de-
scribed using game theory and that this mathematical formalism predicted that
organisms capable of producing apparently indeterminate behavior would be fa-
vored by natural selection.
Imagine, Maynard Smith proposed, a species of animals in which individuals
compete for access to territories that increase the number of young an individual
can produce. Individuals without territories produce a small number of young,
while individuals with territories produce a large number of young. Obviously,
under these conditions it is in the interest of individuals to obtain territories. Now
consider a situation in which there are more individuals than territories. In this
hypothetical species, territories change hands when an animal without a territory
“displays” to an animal with a territory, essentially threatening that individual for
control of the territory. In the hawk-dove game, as this competition has come to be
known, after such a display each animal must make a decision: whether to escalate
the conflict (to fight for the territory) or whether to retreat (give up the territory
without a fight). If one of the animals elects to escalate, behaving as a hawk, and
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34 GLIMCHER

TABLE 1 Payoffs for challenger in the hawk-dove game

Challenger chooses hawk Challenger chooses dove

Defender chooses hawk 50% chance of gaining territory Nothing gained

50% chance of injury
Defender chooses dove Value of territory gained 50% chance of gaining territory

one decides to retreat, behaving as a dove, then the hawk takes the territory. If both
animals elect to be doves, then one of them at random takes the territory. Finally,
Annu. Rev. Psychol. 2005.56:25-56. Downloaded from arjournals.annualreviews.org

if both animals elect to be hawks, then they fight, one sustains injuries that reduce
the number of young that individual can produce, and the other gains the territory.
Table 1 illustrates this simple game as a two-by-two matrix that specifies the costs
and benefits of all possible actions to each player.
by Ball State University on 01/05/09. For personal use only.

What Maynard Smith realized at a mechanistic level was that each of these
values could be expressed in terms of evolutionary fitness, the gain in reproductive
success, that an individual achieves with each outcome. Gaining a territory confers
an increase in fitness, whereas sustaining an injury confers a decrease. Thus, if the
value of a territory is high and the magnitude of injury in a hawk versus hawk fight
is low, then animals that behave as hawks are more fit than those that behave as
doves. Under these conditions, Maynard Smith reasoned, the population will evolve
toward a single pure strategy equilibrium: All animals in the population will always
be hawks. Similarly, if the value of a territory is low and the magnitude of injury is
high, then all animals that behave as doves will produce more offspring, be more
fit, than animals that act as hawks. Under these conditions, the population should
converge on a pure strategy equilibrium of dove. If, however, the value of a territory
is high and the cost of an injury also is relatively high, then an interesting thing
happens. The only reproductively stable strategy for the animal and its offspring
is to behave sometimes as a hawk and sometimes as a dove. To be more specific,
a single dominant and unbeatable strategy emerges in a population playing the
hawk-dove game. The probability that on any given encounter an individual will
choose to behave as a hawk must be equal to the value of a territory divided by
the magnitude of the injury sustained in a hawk versus hawk conflict. Critically,
on each encounter individuals have to behave in an unpredictable fashion, never
allowing their opponent to know whether they will be a hawk or dove2. But across
many such encounters the only stable and unbeatable solution for the population
is for the probability of being a hawk to be equal to the value of a territory divided
by the cost of injury.
This theoretical analysis suggests that, at least from the point of view of
other individuals in this same species, evolution would drive behavior toward

2
Maynard Smith showed mathematically that a population of unpredictable individuals
would dominate a population in which separate individuals were committed at birth to
playing hawk or dove. For details of that proof, see (Smith 1982).
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INDETERMINATE BEHAVIOR 35

unpredictability. As in the game theoretic work of VonNeumann, Morgenstern,

and Nash, the ability to generate apparently unpredictable behavior seems advan-
tageous. One interesting feature of Maynard Smith’s argument, however, is the
mechanism by which this uncertain behavior would be presumed to arise. We
have strong reasons to believe that completely novel behaviors arise, at least in
part, from genetic mutations. Random changes occur in the genomes of these ani-
mals and then selection operates to preserve useful variations in behavioral traits.
Atomic-level fluctuations in DNA molecules, induced by quantum-level forces
like cosmic radiation, produce unpredictable changes in the genetic make-up of a
species. These random changes then influence behavior. We have every reason to
believe that the mechanism by which apparently indeterminate behaviors would
Annu. Rev. Psychol. 2005.56:25-56. Downloaded from arjournals.annualreviews.org

arise would itself be truly indeterminate.

Game theory, whether directed toward the actions of an individual or the evo-
lution of a species, predicts that under some conditions behavior must appear
by Ball State University on 01/05/09. For personal use only.

indeterminate in order for it to be efficient. What implications, if any, does this

have for the determinate scientific method? Does this mean that social scientists
have to abandon Popperian falsifiability? Probably not, for at least two important
reasons. First, the theoretical observation that behavior should appear indetermi-
nate does not mean that behavior does appear indeterminate. Physical constraints
may make it impossible, or unlikely, for mutations to generate behavior that even
appears indeterminate. If this is the case, then at an empirical level we may simply
find that apparently indeterminate behavior does not occur. Second, even if ap-
parently indeterminate behavior were to be observed, this would not require that
the physical generative process for the behavior itself be indeterminate. The psy-
chological system responsible for decision making during strategic games might
operate on totally deterministic grounds. Like a modern digital computer, it may
simply generate an appearance of indeterminacy sufficient to defy prediction by
the opponent. In summary, there may be reasons why fundamental indeterminacies
like those that arise at the quantum level cannot influence the systems that generate
behavior. Either of these two observations would rescue Popperian falsifiability in
its strongest form.

Empirical Measurements of Behavioral Indeterminacy

The theory of games makes it clear that an organism with the ability to produce
apparently indeterminate patterns of behavior would have a selective advantage
over an animal that lacked this ability. Were apparently indeterminate behavior to
have arisen in the evolutionary history of vertebrates, there seems every reason to
believe that this behavioral phenotype would be preserved. Do humans have this
ability? A common answer to this question, based on studies of humans, is no.
Over the course of the last 40 years, a number of psychological studies have
suggested that, perhaps because of some fundamental constraint in the human
nervous system, humans cannot generate behavior that appears indeterminate (for
a review of this literature see Wagenaar 1972). For example, in one of the first of
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36 GLIMCHER

these studies, Bakan (1960) asked humans to simulate the action of a random coin
flip: subjects were asked to make up a sequence of heads and tails that was fully
random in order. When Bakan analyzed the sequences generated by these subjects
they were found to be highly nonrandom despite the instructions that the subjects
received. Bakan found that the subjects tended to overproduce alternations between
heads and tails and to underproduce the occasional long runs of heads or tails that
would be predicted from a truly random process. In sum, the humans behaved in a
fairly determinate fashion, despite their instructions to do otherwise. Since 1960,
dozens of studies have replicated this basic result. When human subjects receive
a verbal instruction to produce a random sequence, they reliably fail. On the basis
of this evidence, many psychologists have concluded that humans lack the ability,
Annu. Rev. Psychol. 2005.56:25-56. Downloaded from arjournals.annualreviews.org

in principle, to generate patterns of behavior that appear indeterminate.

According to game theory, however, environmental conditions should arise in
which apparently indeterminate behavior would be truly beneficial. Organisms
by Ball State University on 01/05/09. For personal use only.

in their natural environment would be reinforced for producing apparently inde-

terminate behavioral patterns under some conditions. Regardless of these human
data, then, can nonhuman animals produce apparently indeterminate behaviors if
they are reinforced for doing so? Blough (1966) was one of the first to ask this
question directly by specifically reinforcing pigeons for producing behavior that
approximated a random process. In that experiment, pigeons were trained to peck
a key in a Skinner box, and the amount of grain that they received after each
peck was contingent upon the length of time that had intervened since the last
peck. The more closely the set of interresponse intervals produced by the pigeon
approximated the output of a random Poisson-like process, the more grain the
bird earned. Blough found that under these conditions the birds quickly adopted
a response strategy that was virtually indistinguishable from the output of a truly
random operator. Figure 1 shows the frequency distribution of interresponse inter-
vals Blough obtained from a single pigeon and, plotted as a solid line, the pattern
of intervals that would be expected from a fully random process. While Blough’s
analysis did not show that the behavior of the pigeons was random by all possible
measures, it did demonstrate that when an apparently indeterminate behavior was
reinforced, pigeons could produce a behavior of this general type. This study was
critical because it provided the first evidence that the ability to produce apparently
indeterminate behavior had arisen in the vertebrate line.
Since that original study, a voluminous literature has examined the ability of
several species of animals to generate apparently indeterminate behavioral se-
quences when they are specifically reinforced for doing so, and tasks more closely
approximating the conditions described by game theorists have also been exam-
ined (see Neuringer 2002 for a review of this literature). Shimp (1967) introduced
one paradigm that has been particularly widely studied. In that paradigm, pigeons
were trained to choose sequentially between left and right response keys for four
responses during each of thousands of trials. The behavior of the pigeons on each
trial thus produced one of 16 possible patterns, for example, left-right-left-left.
The animals then were reinforced for producing the 16 possible patterns with an
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INDETERMINATE BEHAVIOR 37
Annu. Rev. Psychol. 2005.56:25-56. Downloaded from arjournals.annualreviews.org
by Ball State University on 01/05/09. For personal use only.

Figure 1 Mean interresponse times (IRTs) from two replications of three experi-
mental conditions for a single pigeon (Blough 1966). The graph plots the frequency of
each IRT in half-second bins. A fully indeterminate process would produce points that
fall along straight lines in this space. (Reproduced with permission from Journal of
the Experimental Analysis of Behavior, copyright 1966, Society for the Experimental
Analysis of Behavior.)

apparently random frequency distribution. In one important and well-controlled

study, Page & Neuringer (1985) employed a strategy of this type to examine inde-
terminacy in behavior and to see whether the ability of pigeons to produce random
sequences depended specifically upon whether or not they were reinforced for
apparent randomness. In that experiment, pigeons produced long sets of left and
right responses, but under two reinforcement contingencies. Under the first contin-
gency, the animals were specifically reinforced for producing patterns of left and
right responses that had a random-like frequency distribution. Under the second
contingency, the randomness of the emitted frequency distribution was irrelevant
to the reward received. Page & Neuringer found that when reinforcement was con-
tingent on variability, the variability of the pigeons’ responses increased, but when
the level of variability was not reinforced directly, the pigeons adopted much more
determinate response patterns. More specifically, they found that an information
theoretic analysis of the response patterns showed nearly perfect indeterminacy
when, but only when, indeterminacy was reinforced. These results suggest two
interesting conclusions. First, they suggest that the degree of apparent indetermi-
nacy included in behavior is variable. Animals can be more or less indeterminate
based on the requirements of their environment. Second, they suggest that when
Exploring the Variety of Random
Documents with Different Content
that same measure does it owe society a debt. The artist must see for
the people—reveal them to themselves and to each other.”
This is a far cry from “art for art’s sake.” That it is the viewpoint of
an artist of high standing is attested by the early and generous
recognition accorded Miss Eberle’s work from conservative and
radical alike.
Most of her training was received from George Gray Barnard, with
whom she studied for three years. In 1904, she was awarded a bronze
medal at the St. Louis Exposition; the Girl on Roller Skate was
purchased by the Metropolitan Museum in 1907; the Windy
Doorstep was awarded the Helen Foster Barnett prize at the exhibit
of the New York Academy in 1910; her figure of the veiled Salome
was bought by an Italian Art Society in Venice; and she is one of the
ten women who belong to the National Sculpture Society.
Miss Eberle is best known, perhaps, for her dancing figures, and
her depiction of the everyday picturesque life on the lower East Side
in New York, where she lived for years. Her people live for us, and
speak for themselves,—from the placid, necessitous hunt of the Rag
Picker to the tremulous wistfulness of the loving Little Mother; from
the tender feeling of The Bath Hour to the intense, joyous absorption
of the Rag-time dancer and the exultant balance of that flying little
figure on the Roller Skate—and please notice that, characteristically,
there is only one—borrowed, no doubt for a precious three-minute
“coast.”
 THE WINDY DOORSTEP

The qualities which art critics first look for—the sure touch and
line of her modelling, the line composition and massing—are
especially apparent in The Windy Doorstep, in which Miss Eberle
touches the high-water mark of her more objective figures.
Here it will be more interesting to note the steps by which social
values have crept into her work.
First of all, her deep and instinctive love for children, and her
appreciation of human values, led her to select types that until
recently have been almost entirely disregarded.
 To this keen observer and lover of human nature, the many years
of contact with this vivid, arduous East Side life—reinforced and
interpreted by constant reading and thinking—brought an ever-
increasing sense of social interrelation and interdependence. Jane
Addams’ books have, more than anything else, she says, helped to
clarify and mould her vision of the constructive part the sculptor may
play in social readjustment.
This growth of social consciousness has been reflected in her work.

THE LITTLE MOTHER

 Just as The Windy Doorstep, with its fine feeling for the dignity of
everyday homely tasks, outranks the Roller Skater which, she says,
was done in a purely objective spirit, so the White Slave records a
forward step which is a difference of kind even more than degree.

THE RAG PICKER

Here she has turned from her more objective work to the graphic
interpretation of a social menace; and it is here, perhaps, that she
finds herself with surest touch. Her conception of white slavery is as
searching in its indictment, as ruthless, cruel and scourging as the
fact itself. One visitor who saw those haunting figures at the
International Exhibition said afterward:
“I was passing through that room of the exhibit when suddenly I
faced it—I could not go on. I had vaguely realized that this horrible
thing was in the world, but it had never touched me. I sat there for
perhaps an hour, thinking—and thinking—”
This woman was one who has led what is called a “sheltered”
existence, whose instinct would be to turn from any discussion or
writing on this subject. It is this thought-compelling quality in such
work which links it as a social force with, say, the dispassionate but
terrible report of the Chicago Vice Commission, or with Elizabeth
Robins’ My Little Sister.
It is interesting to know that Miss Eberle worked out the
composition for the White Slave four years ago; but the actual work
of modelling was done in the four weeks’ interval between the time
she was invited to send some of her work to last winter’s
International Art Exhibition and its opening. Until then, she had felt
that the time had perhaps not come when such a group would be
received except as an unwelcome effort toward sensationalism. It is
the first of several such interpretative subjects which she has in
mind, and which, if worked out in an equally sincere spirit, should be
big in social significance.
But after all is said, Miss Eberle consistently holds with the many
who believe that the first function of art is not didactic. Much of her
work besides that shown here is conceived in a spirit of sheer joy in
beauty of form and line, and one may go far to find a more
exquisitely modelled figure than that of the utterly submissive,
despairful child-victim of the white slaver. To use her own words:

“It is the beauty that is in the world today that appeals to me—not what may
have existed centuries ago in Greece. Though I love that, too, I will not shut my
eyes to the present and continue to echo the past. No matter how ugly the present
might be, I would rather live in it. After all, ugliness, as well as beauty is in the eyes
of the beholder,—and the present isn’t ugly at all, but full of a wonderful interest,
as a few of us are beginning to find out. We are trying to find new bottles for new
wine—Greek vases are about worn out.”
 THE BATH HOUR

And so we may, as she asks, leave her work to make its own
eloquent plea for what Emerson calls “the eternal picture that nature
paints in the street, with moving men and children, beggars and fine
ladies, ... capped and based by heaven, earth and sea.”
 “MY LITTLE SISTER”

HARRIET BURTON LAIDLAW

[Mrs. Laidlaw is chairman of Manhattan Borough of the
Woman Suffrage Party. Her special interest in the subject of this
review is due, among other things, to her friendship with Rose
Livingston, the rescue worker in Chinatown whose unique
experience gives her understanding sympathy with unfortunate
girls, and who has suffered persecution, unchecked by the police,
on account of her revelations in regard to the white slave trade.—
Ed.]

In My Little Sister[10] Elizabeth Robins has let us hear a great cry out
of the depths—an actual human cry.
True, many snug, comfortable people, shaken for a moment out of
their apathy, dismiss it all carelessly, even contemptuously. Many
who are gripped to the heart as they read brush away the tears with a
sign of relief and say, “Well, it’s only a story.” But what a “story!”
It is told in a tense, staccato style which hurries the reader on—on
even through the idealistic descriptions of the English country and
the stretching moor, across which, in its little garden planted by
loving hands, stands the “dear home” of the officer’s gentle widow
and her two daughters, the Elder Sister in whose words the story is
told and whose name is never mentioned, and the Little Sister,
Bettina, all golden and bright, a creature so touchingly exquisite, so
like a flower.
Through the sweet, lightly sketched scenes of the childhood,
girlhood and dawning womanhood of these two fair young creatures
there is woven a dark thread of fear, dread, tragedy. Some experience
of brutality this dainty, tenderly devoted mother has had, an
experience which is never more than darkly hinted at. She forgets the
terror of it only in those happy memories that cluster about her
lover-husband. His body was brought home to her from a fatal tiger
hunt. Her only consolation in his loss, she tells the Elder Sister, was
that she knew that there are worse dangers than those of the jungle!
She had the joy of knowing that he died while all was “bright and
untarnished.”
The narrative hastens on, always with a hovering sense of doom.
The innocent, tender love stories of the sisters develop. They are but
seventeen and nineteen, but they are women. It is impossible to do
justice to the telling of this story, so we will pass to the main event,
the invitation of the London aunt, the failing health and fortune of
the mother, which nerves her to accept an offer that will separate her
from her children, and the almost happy bustle of preparation for the
first visit from home, and for that wonder of wonders—a London
season! The sinister part played by Madame Aurore, the little
London dressmaker who, while working for them in their country
home lays the plans that are to decoy them to their ruin in London, is
worked out with sure, rapid touch. Then comes the arrival in
London, the meeting of the girls by the pseudo “aunt,” who has been
dressed for the part by the aid of a photograph sent on to the
procurers by Madame Aurore. Then follows the terrible scene in the
house with the barred windows and the strange overwhelming scent,
the escape of the Elder Sister, the heart-breaking search for the
younger—the madness, the despair! One of the most pathetic things
in the book is the heart-broken young lover who traces Bettina from
London to “their house in Paris”—thence from one place to another
“always too late.” The merciful death of the mother, and the fixed
conviction, the subliminal inspiration, that comes toward the end to
the tortured brain of the elder sister that Bettina has found her
release in death, alleviate and spiritualize the misery of it all.
Now as to the truth of this tremendous story. I suppose that having
put the very soul of truth and reality into the narrative itself, having
written it from the very depths of her own pitying heart, the author
has not reckoned with the callous incredulity which the book often
meets, or she might well have added an introductory note in regard
to the actual facts in this case from real life. The story, she told me
subsequent to its publication, was absolutely true, but much softened
at many points. In the true story, for instance, the mother is not
dead, but is in a state of dementia on the continent, whither the
family have moved, being unable to endure England and its
memories.
Moreover, although Miss Robins’ experience in working for
women and the woman’s cause in England had brought to her
knowledge many life histories more unspeakable than this, after she
had finished this book she said to herself, “Now here is a story that I
know to be absolutely true, but how valuable is it to give to the world
an individual instance of this kind; how far can it be generalized?” To
test this she went, with that sincerity of spirit that characterizes all
she does, to a noted police justice in London and laid the story before
him, asking him what he would do if a person came to him with such
a story. He answered sadly, with no show of being at all roused by
anything unusual, that he would begin to take evidence immediately
and see what he could do about it. He added that the story was really
a commonplace.
Such is the reticence of the English press, and such is the stern
family pride concerning the “blot on the scutcheon,” and all those
traditions which grow out of the double moral standard whereby the
escapade of the son of the house can even be repeated lightly while
the slightest shadow on the name of the daughter is an unspeakable
disgrace, that no one knows how many tragedies of this kind are
hidden from sight in the records of numerous respectable families.
To satisfy herself more completely that the story was susceptible of
extensive application, Miss Robins took it to several London police
inspectors. No one for a moment stopped to doubt or question the
facts. The most horrible stories that the human mind can conceive
are old stories to them. When the Home Secretary was asked a
question on this subject in the House of Commons only a few weeks
ago, he answered that in London city alone, to his knowledge, there
had been reported fifty-three girls lost and never heard of within a
few months. Such facts as this explain the tragic intensity with which
the book is written.
Danger! Danger! Danger! That is the dominant note that sounds
throughout the narrative. How pathetically the mother’s poignant
fears contrast with her ineffectiveness. Ever this haunting danger,
whether mother and daughters are walking in the sunset, or planting
in the garden, or sitting by the hearth. What a wonderful picture
Miss Robins gives of the mother’s desperate dependence on the four
walls of her home—“Soon home, now, little girl, soon safe in our dear
home.” The danger signal of the night-bird’s note is introduced with
inimitable art—a subtle suggestion, even in those early days, of the
gray hawk whose shadow hovers over bright young lives.
The unutterable sadness of it all and the stern warning to mothers
that children’s homes are not just in four walls, but are in towns and
cities and nations! How utterly ineffectual seem an individual
mother’s effort for the safety of her child. How evident is it that a
mother’s care must have back of it power—power in council and
legislative hall. How strongly the lack of social sympathy is brought
out; the mother’s indifference to the great crying needs of the world.
This mother’s “place was the home,” and to what did all her negative
efforts avail in shutting the danger away from her cherished
daughters in a nation, in a world, which holds a traffic system of such
Machiavellian adroitness, a system which can afford, so great are its
profits, to reach into the inner recesses of a home, to work with
endless patience and resourcefulness and which can enlist on its side
such power that even the London police, perhaps the least corrupt in
the world, can answer evasively to the frantic cry, “Do you know such
a house?”—“We have a great many on the list, but not many such as
you describe;” and follow this statement with the maddening
inactivity which Miss Robins describes with vivid accuracy.
Her book is a terrific arraignment of the conditions which make
such a tragedy possible. Respectability and indifference are
personified scathingly in the monumental aunt, deaf to the world
voices of agony. All society is arraigned as the Elder Sister storms at
her aunt, “sitting massive, calm, with a power of inert resistance.”
Her bewildered answer to the mad cry for help,—“It isn’t possible,
this is England,”—sounds strangely natural to us. It is the burden of
so many recent New York editorials, “such things don’t happen”!—
and that in our land where the record of the Chicago Immigration
League tells of 1,700 girls between the railroad terminals of New
York and Chicago alone, reported lost in one year. Thus wails the
Elder Sister,—“So old and unbelieving, I felt she had looked on
unmoved at evil since the world began.... She rose, O! but slowly;—
slow, stiff and ponderous. I felt in her all the heaviness of
acquiescence since time began.”
Thus is the unbelievable apathy of society pictured! Thus Miss
Robins touches lightly, pitifully on the problem of a girl’s handicap in
the lack of preparation for life. What a picture she gives of the
sheltered girl—“Such a little thing, my not knowing how to
telephone, yet it might cost my mother her life.” Again this motive is
sounded when the daughter is begging her mother for knowledge
about her experiences, and the great gray danger—“It is not the kind
of thing you need ever know,” answers the mother with fatuous
finality.
Nor does she make our heart bleed for girlhood alone, but for
manhood, as that blood-curdling conversation, unparalleled in
literature, is gasped out between the Man and the Elder Sister as they
crouch in the shadow in that ghastly room in what he himself tells
her is “one of the most terrible houses in Europe.” Here we see
manhood disfigured, draggled beyond recognition, sitting in the dust
and ashes of a charnel house. How sodden the words fall from his
lips as depraved and perverted manhood defends itself. “This is a
commonplace in the world, in every capital of every nation on earth.
Bishops, old ladies, imagine you could alter these things.” “Human
nature—human nature,” muses the Elder Sister, “like the tolling of a
muffled bell.” Thus are the wickedest, most deadly of the world’s
platitudes on this subject uttered in this glaring scene of
abnormality, so that automatically they are given the lie.
The unbelieving will say, “But how is it possible that we do not
hear oftener of such cases?” Two elements especially conspire to
suppress such knowledge, both based on a false attitude towards
women. First the double standard of morals which holds men’s
purity so much lower than woman’s; second, the unjust attitude
towards women reflected in the press which is more likely to seize
upon a disappearance story in such a way as to make it reflect upon
the girls’ character rather than to acknowledge it a possible case of
abduction or white slavery.
As I write, Miss Robins sends me letters which she has recently
received from people interested in her book. One speaking of the
“awful traffic that is going on” tells how two girls, daughters of a
clergyman, have disappeared on their journey home from school and
have never been heard of since.
A letter from Miss Robins herself adds the following to her
remarks on her book. “Of all the official people I consulted, not one
of these experts doubted my story, and all had known similar cases.
We do not need to be told that the people to whom these things
happen, if they are refined and sensitive, are not eager to make
known their ruin. The natural impulse is to cover the horror from
every eye, even to deny it.”
“If you will consult the findings of your own commissions you will
see that in America ‘no girl of any class is safe.’”
I myself know at first hand an appalling number of cases—wives,
women of standing, college girls on journeys, young girls on their
way to boarding school, and working girls trapped and sold. In Rose
Livingston’s mail in the last few weeks have come heartrending
letters from parents for help in finding their lost daughters. A letter I
read yesterday from a doctor’s wife in Indiana was so terrible in the
lingering pathos with which the mother dwelt on the beauty and
sweetness of this lost daughter that I handed it back without reading
to the end.
In connection with Miss Robins’ letter, I offer the following
quotation from a paper by Stanley Finch of the Federal Department
of Justice where he speaks of the operation of the Federal White
Slave Traffic Act. “The number of complaints and prosecutions is
rapidly increasing.... Such crimes are more numerous than was first
believed possible.... It has become evident that thousands of
people[11] in practically all parts of the country were violating the
law.... [We found] girls were being transported in such interstate and
foreign commerce solely for the purpose of prostitution and were
being treated as mere articles of merchandise for the profit of those
who handled them and who were willing for the profit involved to
sacrifice both the bodies and the souls of their victims.... As we
extend this work [of federal prosecution] throughout the country the
awful interstate trade in women and girls for immoral purposes
which for years has been going on almost without let or hindrance,
has now become a very dangerous occupation.... The practices of
people engaged in the white slave traffic involves in a considerable
number of cases the actual physical detention of women and girls
against their will in this vilest form of involuntary servitude.”
Would that there were space to reinforce these remarks by New
York newspaper headlines dealing with such cases for the month of
March 15 to April 15.
How many fathers and mothers, brothers and sisters, friends and
lovers throughout this country, have lived through the agony
endured by the Elder Sister and the lover in Miss Robins’ book? That
which had been their greatest care and burden, the frail health of
Bettina, becomes the Elder Sister’s one comfort and hope. How many
a bereft father and mother today not only cling in anguish to that
hope, but desperately refuse to believe but that their girl is dead! And
with reliable students estimating the life of the white slave in this
country at five years, we know that many a flower-like highly
organized Little Sister has met mercifully her release in a few weeks
—a few months.
To those who know how considerable a factor in the whole
problem of the white slave traffic is the girl who is taken, not the girl
who goes, the girl who is under compulsion, not the girl who stays,
this book is a great contribution. The Survey printed some time
since a poster that has been used in this country from ocean to
ocean: “Danger! Mothers, beware! 60,000 innocent girls wanted to
take the place of 60,000 white slaves who will die this year in the
United States.” If My Little Sister will only make the truth of this
warning more real, more individual more poignant, then, in the pain-
soothing words that close this book, “She will not have suffered in
vain, and others will thank her too.”
Such a book is not merely a literary production, an exquisite work
of art: it is a high, sincere human service. Front a literary point of
view it is a great book. One is reminded of the Aeschylean definition
of tragedy: “That which purifies the heart through pity and terror.”
But this book not only reaches great tragic and dramatic heights, but
its subtle art is such that it blends with the tragedy in an almost eerie
way a lyric chord which echoes throughout, an unbroken strain of
hope and pity, of the essential dignity and sanity and rightness of life.
 MOTHERHOOD

Isabel Kimball Whiting

I see them come crowding, crowding,

Children of want and pain,
Dark sorrow their eyes enshrouding
Where joy’s touch should have lain.

They stand in silence beseeching,

Gaunt faces lifted up
And wan little hands outreaching
For Love’s forbidden cup.

Their hearts are restless with yearning.

The hearts of my own are stilled,
Their lips are parched and burning
The cups of my own are filled!

I cry in love unsatisfied

For these without the fold,
My mother’s arms are open wide
These weary ones to hold.

What though my arms are open wide,

Only mine own lie near.
Without still stand those long denied,
Compassed in want and fear.

Bowed with the crown of Motherhood,

I seek that Shepherd of old;
“How can mine own receive the good
With some left out of the fold?”
 IN THE HANDS OF THE POLICE

A PLEA FOR A NEW LEGISLATURE TO MAKE AND ENFORCE

LAWS AGAINST VICE

A. LEO WEIL
[Mr. Weil, one of the best known attorneys of western
Pennsylvania, is known outside of his profession as the militant
President of the Pittsburgh Voters’ League, the organization
which exposed the graft scandals of three years ago, sent
councilmen and bankers to jail and brought new and invigorating
spirit into Pittsburgh municipal life. Last Year the Voters’ League
successfully brought charges against the head of the Department
of Public Safety, and provoked a situation which led to the
creation of the Pittsburgh Morals Commission, a semi-official
body without governmental authority, and the cleaning-up of the
big numbered district.
It is on the basis of this experience that Mr. Weil reaches the
conclusions here set forth. He was a witness before the Wagner
commission which has recommended legislation now pending in
New York (with every prospect of passage) that will create a
separate public welfare department, largely divorce it from the
city administration and entrust its commissioners, appointed by
the mayor, with powers with respect to vice somewhat similar to
those held by the Board of Health with respect to sanitary
conditions.—Ed.]

The police, a body of men selected primarily to preserve order,

protect life, prevent crime, apprehend the criminal, and perform
other administrative duties—men perhaps wholly unfitted as a body
for anything else—have been expected to solve, by legislation,
problems which have confounded the wisest from the time whereof
the memory of man runneth not to the contrary.
That the police have failed could not be otherwise.
That they have aggravated the evil was to be expected.
That ultimately legislation must be placed in competent, qualified
hands, must be apparent.
That it will require the greatest minds, the best thought, the
highest statesmanship, and almost a divine perception, to apprehend
and deal with those complex, involved, intricate questions having to
do with the passions of men and the strongest laws of nature, urging
defiance of human laws, seems to be axiomatic.
Nevertheless, on the subject of vice as generally understood in our
cities, the police are expected not only to administer the laws, but to
make them; not only to enforce enactments, but to frame them. And
herein lies, in my judgment, the root of the evil in present day
conditions which has brought our police into disrepute.
All countries, from the most despotic to the most liberal, from the
most conservative to the most radical, have left this question of the
social evil to the police, and have not dared, through their legislative
bodies, to thoroughly consider and pass laws to eradicate it. Such
meager legislation as has been passed has been ignored in the
congested centers of population, and generally such legislation has
not been supported by the public sentiment of those communities.
Legislation has been generally of a merely prohibitive character,
ignoring existing conditions, together with the army of human beings
living upon vice, for whom some provision must be made.
Even if the legislatures could be induced to take up this question
and pass general laws, it would necessarily follow that such laws in
their generality must look to the future, rather than to the present,
and existing conditions would have to be left to some other authority.
In every large city there are thousands engaged in prostitution.
They can neither be exterminated by the fiat of law nor reformed by
the passage of proclamations. Public morals, like private morals, can
be improved only gradually. It is not so much the theoretical
question of to be or not to be, but the practical situation summed up
in such phrases as: It is—it will for a time remain. The future—what
is it to be?
Would it not be wise to ask the legislature to pass a law requiring
every city to appoint a body of men to draft, and from time to time
revise, a code of laws to regulate, suppress, exterminate, and
generally to deal with, this problem of prostitution? To it must be
given authority. It must be vested with legislative power. It should
have its own employes—call it a morals police force, if you wilt—to
carry into effect the rules, regulations and laws passed by this
commission. That men could be obtained to act upon such a
commission, public-spirited, representative students of social
questions, I fully believe. No one would accept a place on such a
board who was not interested in the welfare of his community. The
responsibilities would be fearful; the criticism would be severe. No
one without a blameless life would dare to act. Whether or not
ultimately the regular police force could be used by such a
commission would be a question of policy. At any rate, for the
present, I believe it would be unwise.
Such a commission would collect data of inestimable value, data
that was reliable, and of which there is now none worth while. Such a
commission would doubtless make mistake after mistake, perhaps
would depart from all its preconceived notions at the start.
Nevertheless, it is reasonable to assume that step by step it would
gradually and slowly work out a solution. Looking the country over,
we see that voluntary organizations have accomplished temporary
good, but their effort has never been persistent, continuous,
permanent, authoritative, legal.
The adoption of the plan suggested would take out of the hands of
those wholly unfitted for so great an undertaking this momentous
question, and place it in the hands of others who we would have the
right to assume would ultimately be found to deal with it, so far as
human intelligence, study, observation and experiment can enable
and qualify men to deal with it.
Another and a greatly to be desired result would also be
accomplished. It would remove from the police that which today
contaminates the organization, so that in every city they have been
brought under a suspicion unfortunately too frequently deserved, an
opprobrium too justly applied.
 The casting of this burden upon the police, more than any other
cause, sows the seed of corruption, furnishes the opportunity for
profit, brings into alliance the law officers and the law-breakers,
disgraces the police force, and keeps off of it many men who would
otherwise be glad to serve their communities in a position that
should command the respect and consideration of the people.
This police burden has in turn been the biggest rock upon which
self-government has capsized in our American cities.
 THE BELGIAN STRIKE

[The mass strike of the Belgian workers was carried to an issue

last week. The motion of the Liberal member which brought the
strike to a close, while it does not specifically grant the reform
asked for by the Socialists, is considered such a lien on
government action as to amount to a victory for them. These
interpretations of the larger significance of the strike from
different angles were, of course, written while the struggle was
going on.
Mr. Walling, who is a member of the radical wing of the
Socialist party, is a student of international Socialism, his most
recent book being The Larger Aspects of Socialism. He was in
Russia during the Revolution, and knew intimately the details of
the great Russian general strike of 1905.—Ed.]
 I.
GRAHAM TAYLOR
The nation-wide strike for manhood suffrage in Belgium which
paralyzed the industries and almost suspended the normal life of the
people was foreshadowed in a spectacular demonstration in Brussels
two years ago.
On August 15, 1911, the streets of that gay, medieval capital
witnessed scenes which every American who looked on knew were
making history. Over 60,000 men, from larger and smaller places
throughout Belgium, took a day off without wages and paid their way
to the capital of their country, in order to voice their protest against
the unjust inequalities of the suffrage. The show of force by the
extraordinary police and military precautions betrayed the furtive
apprehension of both the municipal and national governments as to
what might happen. With no sign of timidity or intimation of being
overawed, this vast industrial army marched ten abreast for hours—
silent, grim, determined, united, unarmed—between long files of
armed soldiery which lined the curbs, and past stronger detachments
of all arms of the service massed at strategic centers.
The great procession assembled at the Socialist headquarters, a
large and impressive building bearing the significant name Maison
de Peuple, the House of the People. The permanent background of
the stage in the assembly hall of that building is a colossal head of
Jesus of Nazareth, the reverent work of a Belgian Socialist. This
House of the People is the most practical expression, or perhaps
demonstration, of the co-operative commonwealth in miniature,
which is to be found anywhere in the world. Starting with a sack of
potatoes and a bag of flour, these wage-workers in ten years erected a
building costing $250,000. Of this sum, which was loaned by the
national bank, they had then paid $100,000 and had assets worth
three times as much as the balance due on the mortgage, which they
continue to reduce by annual payments.
In this four-story semi-circular building, at one of the principal
business centers, ample accomodations are provided for a great
variety of practical agencies. A café, which paid a profit of $2,400 in
three months, shares the front of the ground floor with a large co-
operative department store, where dry goods, house furnishings,
clothing, meats, groceries, butter and milk, hats, hosiery and shoes
are sold. A bakery, with a capacity of 125,000 loaves of bread a week;
a coal depot, with twenty-nine delivery carts; a laundry, and a
clothing manufactory are among the business enterprises conducted
here.
The 19,000 co-operating families receive as their share of the
profits 12 per cent of the money they pay for bread, 6 per cent of
what their groceries cost them and 5 per cent of the purchase price of
their clothing. Among the protective features are an employment
bureau for men and women, a pharmacy and a corps of thirteen
physicians rendering free service to all members of one year’s
standing, and a sick benefit society with 8,000 members. Singing
and ethical classes are maintained for children and a well-trained
orchestra and choral club for adults. Small halls adequately provide
for the meetings of the trade sections, and a great auditorium,
seating 2,436 persons, rallies the festival gatherings and supplies
room for political mass meetings.
From this national center the procession of mid August took up its
line of march, carrying banners which took the keynote of their
inscriptions from the following figures emblazoned everywhere:
993,070 have 1 vote;
395,866 have 2 votes;
704,549 voters, having two, three or four votes, cast 1,717,871
votes, a majority of 88,523 over those having one vote;
One man one vote!
In Belgium a man over twenty-five years of age gets an extra vote if
he owns property. He is granted another vote if he has a university
diploma. He casts a fourth vote if he is over thirty-five years of age, is
the father of a family and pays taxes on more than a certain amount
of property. The majority of the industrial population thus have only
one vote, while the rural, well-to-do and richer people outvote each
wage earner by two, three or even four votes. The rural population
thus controls the city industrial population and the church is charged
by the Socialists with controlling the rural vote.
 Against this rule of the minority, this great demonstration of 1911
was a protest. But to the onlooker from abroad it then seemed to be a
patriotic proclamation of Belgium’s one great hope of national
evolution without revolution. The primary cause of the movement
which has culminated in the present national strike was the defeat of
the liberal and Socialist coalition in parliament by a combination of
the government and clerical forces in the elections of 1912. The
Socialist congress summoned to meet the issue brought to a crisis by
that event decided upon a general strike as a last resort if all other
means of obtaining manhood suffrage failed. But before resorting to
that measure a general suffrage bill was introduced into parliament
by the Socialists and supported by the liberals. As serious
consideration of it was refused by the clerical and government
authorities, a general strike was voted on April 14.
Whatever the full effect of this national political strike may be,
those who are the keenest observers concede that the making of
history is in the movement of Belgian labor for one man one vote
suffrage.
Certain it is that a movement of the people capable of maintaining
and increasing for so many years a labor vote in parliament until it
numbers more than one-third of the total must be reckoned with. If
now the tolerance of this Belgian Socialist Party toward those who
honestly oppose its principles and methods, at this supreme crisis in
its history and the national development, grows with its strength and
equals its determination, it will improve the greatest opportunity the
Socialist cause has ever had in the sphere of practical politics to
demonstrate and promote its co-operative commonwealth.
 II.
WILLIAM ENGLISH WALLING
If the Belgian strike is a world-event of the first magnitude for the
general public, it has a still greater significance for the world’s ten or
twenty million Socialists. The two wings of the Socialist movement
are equally interested; the reformers and conciliators, because the
strike aims at a purely political reform, and involves co-operation
with a part of the capitalists, both in order to win success now and in
order to get immediate use out of the suffrage after it is won; the
revolutionists and advocates of class struggle because the strike
forces a large loss on many unwilling employers and gives training
for later and more aggressive strikes for the purpose of raising wages,
cutting down profits, and paving the way to social revolution.
These essential facts are being widely understood. Take, for
example, the following editorial paragraphs from the influential and
progressive, but by no means radical, Chicago Tribune:
“For years the thinkers of the movement in Europe were building
up theories about the ‘political mass strike.’ These theories have
now been put into practice in Belgium with remarkable precision.
“The strike in Belgium is not a precipitated strike. For months
the Socialists of that country have been making preparations for it.
They have been collecting money, storing provisions, and, what is
even more important, educating the workingmen to their theories,
training them to respond to the strike call like drilled soldiers. It is
not an emotional, not an impulsive strike, therefore, but a coolly
thought out, shrewdly calculated battle....
“As a result of this careful planning and training by the leaders
400,000 workingmen responded to Socialist colors with the
precision of a trained army. The strike is both a battle and military
review. The Socialists now have a clear and adequate view of their
strength and numbers. The issue involved in this strike, which is
uniform as opposed to plural voting, may be lost. The strike may
even be called off by the Socialists themselves after a week or so if
the government does not yield by that time. But the advantages
 which Socialism has gained by this census of its army is
incalculable. It knows how much of the Belgian public is behind it
and to what extent. This time the general strike is used to combat a
single abuse of the government. In the future the same working
masses may be directed against the present régime as a whole. It is
a strike now. It may be a civil war or revolution the next time.”
But while the capitalistic Tribune is so sympathetic and optimistic,
the leading labor union paper of this country, the United Mine
Workers’ Journal has been highly sceptical and pessimistic:
“The strikers say they are ready and have the means to hold out
six weeks.
“What if the other class should elect to hold out for twelve weeks;
that is, if the strike should become a ‘lock-out?’
“There is no doubt but that the class against whom they are
striking, the propertied class, who hold the unfair political
advantage over them, could store more per member than the
workers.
“We wish our brothers every success. Hope they will obtain what
is undoubtedly theirs by right. But, in our opinion, unless the strike
results in a test of force instead of passive endurance (and this
same is more than probable) the workers’ needs will drive them
back to their tasks before those who have been enriched by their
labor will consent to give up the political advantages on which their
economic advantages are based.”
Why this remarkable reversal of the opinions that might have been
expected? Why have some of the richest Liberals in Belgium
supported the strike?
The Belgians are striking for a right the French obtained in 1876—
equal suffrage. Yet who are the chief beneficiaries of the political
democracy of France? The condition of the laborers is about the
same as in Belgium. The only difference is that in France the
industrial and urban capitalists now hold the balance of power
between the laborers on the one hand and the reactionary agrarians,
landlords and employing peasants on the other; while in Belgium the
latter classes, which control the Catholic Party, have a Parliamentary
majority over the laborers and urban middle classes combined. The
Belgium Liberals then have had everything to win by a strike, which
aims at establishing the French situation in that country. As for the
Socialists, equal suffrage would undoubtedly have the effect of
driving the Liberals and Catholics together into the same
government, as in Germany, thus making the Socialists the
opposition party. It would also result ultimately in whatever social
and labor reforms the Liberals might feel it to their interest to grant.
But this is all.
This is why thoughtful Socialists the world over, including several
of the Belgian leaders (Vandervelde, Huysmans, De Brouckère and
Bertrand) have been so dubious about the strike. They have seen that
the burden of the contest will fall about equally on employers and
employes. But the Liberal employers are playing for a splendid stake
—Belgium; while the Socialists are playing only for such incidental
and secondary benefits as will accrue to them from Liberal control of
the country. Naturally they have had far more dread of the costs and
risks which the strike involves.
But whether the strike was to be lost or won it has been clear that
it would interest the world’s millions of Socialists more deeply than
ever in the possibilities and the limitations of the general strike.
All Socialists favor the political general strike, most Socialists favor
the general strike against war—including the conservatives like Keir
Hardie and Jean Jaurès. The moderate Swedes had an economic
general strike a few years ago and this form is favored also by the
majority of French and Italian labor unionists. And now the British
unionists are voting on the question whether they will all quit work
each day after eight hours—which would mean a general lock-out, or
practically a general strike.
Whether the Belgians win or lose will not affect the momentum of
the movement. If they win, general strikes for a few years will take a
predominantly political character, and we shall see the general
political strike resumed in Hungary within a few months and
doubtless declared in Prussia before many years. If they lose these
British, French and Italian movements toward economic general
strikes will have the field.
The historian of the future when writing of our generation will
have to give a central position—perhaps the central position—to the
general strike.
 1. See The Survey for October 5, 1912.

2. It is now reported that the opponents of the law, which goes into effect
August 1, have started an effort to secure the 19,535 signatures necessary for a
petition to have the law submitted to a referendum vote.

3. Drawn from a case record of the New York Charity Organization Society.

4. Number based not on total number of cases but on scholarship, conduct

and attendance cases, respectively.

5. Most of the quotations here given are taken from Songs from the Ghetto. By
Morris Rosenfeld. Translated into prose by Leo Wiener. Copeland & Day, Boston.
1898. Price $1.00.

6. Rosenfeld’s poetry in this respect is the antithesis of that of Arturo

Giovannitti. See The Survey of November 2, 1912.

7. Pendulum.

8. I work and work and work—without end; I am busy and busy and busy at all
times.

9. The Counting—the forty-nine days of the Feast of the Seven Weeks.

10. My Little Sister. By Elizabeth Robins. Dodd, Mead and Co. 344 pp. Price
$1.25. By mail of The Survey $1.37.

11. Of 478 defendants, fifty-three women are involved or implicated. Of those,

eighteen are found not guilty, are dismissed, or cases are pending.
 TRANSCRIBER’S NOTES
Typos fixed; non-standard spelling and dialect
retained.
Used numbers for footnotes, placing them all at
the end of the last chapter.
*** END OF THE PROJECT GUTENBERG EBOOK THE SURVEY,
VOLUME 30, NUMBER 5, MAY 3, 1913 ***

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