Neuropsychology

1997, Vol. ll.No.1, 104-113

In the public domain

Emotional Perception and Memory in Amnesia

Stephan B. Hamann
University of California, San Diego

Larry Cahill
University of California, Irvine

Larry R. Squire
Veterans Affairs Medical Center and University of California, San Diego
The authors examined whether perception of emotional stimuli is normal in amnesia and
whether emotional arousal has the same enhancing effect on memory in amnesic patients as it
has in healthy controls. Forty standardized color pictures were presented while participants
rated each picture according to emotional intensity (arousal) and pleasantness (valence). An
immediate free-recall test was given for the pictures, followed by a yes-no recognition test.
Arousal and valence ratings were highly similar among the amnesic patients and controls.
Emotional arousal (regardless of valence) enhanced both recall and recognition of the pictures,
and this enhancement was proportional for amnesic patients and controls. Results suggest that
emotional perception and the enhancing effect of emotional arousal on memory are intact in

Many studies have indicated that emotional material is

more memorable than neutral material (Christiansen &
Loftus, 1987; Craik & Blankstein, 1975; Heuer & Reisberg,
1990; Matlin & Stang, 1978). The primary factor underlying
the effect of emotion on memory appears to be emotional
arousal, independent of whether the arousal is pleasant or
unpleasant. In studies of nonhuman animals, emotional
arousal has been demonstrated to affect memory through the
interaction of peripheral adrenergic systems with cholinergic, opioid peptidergic, and GABAergic systems in the
amygdala (McGaugh et al., 1993). The amgydala also has
been implicated in Jear conditioning (Davis, 1994; LeDoux,
1993).
Although emotional arousal also can benefit memory by
its effects on cognition (e.g., increased attention, deeper
elaborative processing), the enhanced memory associated
with physiological arousal is independent of these cognitive
factors and can be dissociated experimentally from them.
Thus, a recent study in humans (Cahill, Prins, Weber, &
McGaugh, 1994), involving memory for an illustrated story,
demonstrated that the memory facilitation associated with

emotional arousal was selectively abolished by propranolol,

a p-adrenergic antagonist. Yet, participant ratings of the
emotional arousal produced by the story were unaffected by
the drug.
Several studies of experimental animals indicate that the
amygdala is involved in the enhancing effects of emotion on
memory (McGaugh et al., 1993). Findings from a patient
with bilateral lesions apparently restricted to the amygdala
(Cahill, Babinsky, Markowitsch, & McGaugh, 1995) are
consistent with evidence from experimental animals. Using
the same emotional memory task as in the study with the
p-adrenergic antagonist (Cahill et al., 1994), this patient
failed to benefit from emotional arousal, suggesting that the
amygdala plays a crucial role in mediating the memory
facilitation effect in humans.
Bradley, Green wald, Petry, and Lang (1992) examined the
effect of arousal (emotional intensity) and valence (pleasantness) on memory in college students, systematically examining the relative influence of each factor on memory. Stimuli
were drawn from the Internationa] Affective Picture System
(IAPS; Lang, Ohman, & Vaitl, 1988), a standardized set of
color slides depicting a wide variety of naturalistic subjects
that spans a wide range of affective reactions. The primary
finding was that the effect of emotion on memory (both
recall and recognition) was mediated by arousal rather than
valence. Specifically, after the effect of emotional arousal
had been partialed out, there was virtually no effect of
stimulus valence (pleasantness) on memory. This study and
others (Lang, Bradley, & Cuthbert, 1990; Russell, 1980)
have validated the dimensional approach to measuring
emotion (arousal and valence). We have adopted the same
dimensional approach in the current study.
Here, we examined whether the effects of emotion on
memory would be observed in patients with amnesia. Little
is known about the effect of emotion on memory in amnesic
patients or even whether the emotional reactions and emotional perceptions of amnesic patients are normal (however,

Stephan B. Hamann, Department of Psychiatry, University of

California, San Diego; Larry Cahill, Center for the Neurobiology
of Learning and Memory, University of California, Irvine; Larry R.
Squire, Veterans Affairs Medical Center, San Diego, and Departments of Psychiatry and Neurosciences, University of California,
San Diego. Stephan B. Hamann is now at the Department of
Psychology, Emory University.
This research was supported by the Medical Research Service of
the Department of Veterans Affairs, National Institute of Mental
Health Grant MH24600, the McKnight Foundation, and the
McDonnell-Pew Center for Cognitive Neuroscience. We thank I.
McGaugh for his helpful comments and N. Champagne and J.
Zouzounis for research assistance.
Correspondence concerning this article should be addressed to
Larry R. Squire, Department of Psychiatry (116A), Veterans Affairs
Medical Center, 3350 La Jolla Village Drive, La Jolla, California
92161. Electronic mail may be sent to lsquire@ucsd.edu.

104

105

PERCEPTION AND MEMORY IN AMNESIA

see Hamann et al., 1996). Reports of emotional responsivity

in the amnesic syndrome have focused almost exclusively
on patients with alcoholic Korsakoff s syndrome (Rapaport,
1961; Talland, 1967). Although these reports concur in
characterizing the emotional responses of patients with
Korsakoffs syndrome as abnormal, they differ in their
descriptions of the patients. Some patients are described as
having reduced reactivity to emotional stimuli (sometimes
referred to as flattened or blunted affect), whereas other
reports characterize other patients as overly irritable and
hyperarousable. Similarly, the few experimental studies of
emotional reactivity in amnesic patients with Korsakoffs
syndrome (Douglas & Wilkinson, 1993; Markowitsch,
Kessler, & Denzler, 1986; Oscar-Berman, Hancock, Mildworf, Hutner, & Weber, 1990) have yielded inconsistent
results. Some researchers have found their reactions normal
(Douglas & Wilkinson, 1993), whereas others have found
them either less reactive (Markowitsch et al., 1986) or more
reactive (Oscar-Berman et al., 1990) than controls. The
interpretation of these studies is complicated because different measures and materials were used to assess reactivity
(e.g., self-reported ratings, skin conductance response, assessment only of negative emotions).
Apart from the issue of emotional perception in amnesia
is the issue of whether the effect of emotional arousal on
memory is normal or abnormal. Anormal effect of emotional
arousal on memory in amnesia would be consistent with two
notions: First, the brain structures that mediate the effect of
emotional arousal on memory (e.g., the amygdala) modulate
declarative memory (McGaugh et al., 1993). Second, these
modulatory brain structures are distinct from the medial
temporal lobe structures important for declarative memory
(Squire & Zola-Morgan, 1991). Alternately, the effect of
emotional arousal on memory may be abnormal in amnesia
because the perception or expression of emotion is itself
impaired in amnesia.
The purpose of the current study was to examine whether
emotional perception is intact or impaired in amnesia and to
determine whether amnesic patients, like normal individuals, exhibit better memory for emotionally arousing material
than neutral material. Perception and memory was assessed
using the standardized IAPS picture stimuli (Lang et al.,
1988) in a group of 9 amnesic patients, 2 postencephalitic
patients who were amnesic but also had damage to the
amygdala, and two groups of controls. Participants rated 40
pictures individually for emotional arousal and valence and
were given a free-recall test for pictures they had viewed,
followed by a yesno recognition test.
The first group of controls (n = 12) was administered the
same test procedure as the amnesic patients (n = 9) and the
postencephalitic patients (E. P. and G. T). The second group
of controls (control-delay; n = 12) was given a longer
interval between the end of the study phase and the memory
tests. The purpose of the delay was to bring the control-delay
group's level of memory performance into the amnesic
patients' range. This condition was important because it was
possible that the effect of emotional arousal on memory
would differ depending simply on whether the overall level

of memory was high or low (e.g., in the control and the

amnesic groups, respectively).
The first question of interest was whether the ratings of
amnesic patients for the emotional stimuli would be normal
or abnormal. The second question was whether the effect of
emotional arousal on memory in these patients would be
normal or abnormal. Another question concerned whether
the patients with Korsakoffs syndrome would exhibit abnormal emotional reactions, as previous reports have described,
and whether their reactions would differ from those of other
amnesic patients. This was of interest partly because patients
with Korsakoffs syndrome sometimes have frontal lobe
dysfunction along with their memory deficits, and frontal
lobe dysfunction has been linked to emotional deficits
(Stuss, Gow, & Hetherington, 1992). Finally, the emotional
reactions and the effects of arousal on memory were of
particular interest in the case of the 2 postencephalitic
amnesic patients with amygdala damage because of the links
among amygdala, emotion, and emotional memory.

Method
Participants
Amnesic patients. Nine amnesic patients (6 men and 3 women)
participated in this study. Four patients had Korsakoffs syndrome.
All 4 had participated in quantitative magnetic resonance imaging
(MRI), which demonstrated reductions in the volume of the
mammillary nuclei (for R. C., J. W., and P. N., Squire, Amaral, &
Press, 1990; for N. F., unpublished observations). A 5th patient (M.
G.) sustained a bilateral media] thalamic infarction, which was
confirmed by MRI. Of the remaining 4 patients, 3 had bilateral
reduction in the size of the hippocampal formation confirmed by
MRI (for P. H., Squire et al., 1990; for L. J. and H. W., unpublished
observations). Patient P. H. had a 6-year history of 1- to 2-min
attacks (of possible epileptic origin) in association with gastric
symptoms and transient memory impairment. In 1989 he suffered a
series of small attacks that resulted in marked and persisting
memory impairment. Patient L. J. became amnesic gradually
during 1988 and 1989 without any known precipitating event. Her
memory impairment has remained stable since that time. Patient
H. W. became amnesic gradually beginning in 1987 and has
remained stable for about 7 years. Slight declines in other cognitive
functions were noted in 1994 and 1995, leading to a diagnosis of
probable Alzheimer's disease. The 4th patient (A. B.) was unable to
participate in MRI studies, but the etiology of his amnesia (anoxia)
is consistent with hippocampal damage. All 9 patients were
well-characterized neuropsychologically (see Tables 1 and 2).
The patients' mean age was 63.4 years at the beginning of the
study, and they had an average of 13.6 years of education.
Immediate and delayed (12-min) recall of a short prose passage
averaged 4.0 and 0 segments, respectively (Gilbert, Levee, &
Catalano, 1968; maximum number of segments = 21). Scores on
other memory tests are shown in Tables 1 and 2. The mean score on
the Dementia Rating Scale was 131.8 (Mattis, 1976; maximum
score = 144), with most points being lost from tile Memory
subscale (mean points lost = 7.3). The mean score for the Boston
Naming Test was 56.0 (Kaplan, Goodglass, & Weintraub, 1983;
maximum score = 60). The number of categories completed on the
Wisconsin Card Sorting Test (Heaton, 1995) for Patients R.C., N.F.,
P. N., J. W., M. G., H. W., A. B., P. H., and L. J. were 4,3,3,5,4,2,
5, 5, and 6, respectively. Scores for controls on these tests can be

106

HAMANN, CAHUJL, AND SQUIRE

Table 1
Characteristics of Amnesic Patients

Patient

Age
(years)

R.C.
N.F.
P.N.
J.W.
M.G.
H.W.
A.B.
P.H.
L.J.

M
G.T.
E.P.

76
57
65
56
60
77
55
70
55

Lesion
Dien
Dien
Dien
Dien
Dien

HF
HF
HF
HF

63.4

59
73

WMS-R

WAIS-R

HF+
HF+

66.0

Attention

IQ

Verbal

106
94
99
98
111
109
104
115
98

115
91
81
104
113
97
87
117
105

76
62
77
65
89
84
62
67
83

103.8

101.1

73.9

92
103

120
94

57
57

107.0

57.0

97.5

Visual

General

97
73
73
70
84
102
72
83
60

80
53
67
57
86
89
54
70
69

79.3

<50
82
66.0

69.4

<50
61
55.5

Delay

72
<50
53
57
63
54
<50
57
<50
56.2

<50
56
53.0

Note. The WAIS-R and the WMS-R indexes yield a mean score of 100 in the normal population,
with a standard deviation of 15. The WMS-R does not provide scores for those who score below 50.
Therefore, the six scores below 50 were scored as 50 for calculating a group mean. WAIS-R =
Wechsler Adult Intelligence ScaleRevised; WMS-R = Wechsler Memory Scale-Revised. Dien =
diencephalon; HF = hippocampal formation; HF+ = hippocampal formation and additional brain
structures.
found elsewhere (Janowsky, Shimamura, & Squire, 1989; Squire et
al., 1990).
Patients E. P. and G. T. Two additional patients (E. P. and G. T.,
both postencephalitic patients) also were tested. These 2 patients
are considered separately from the amnesic group both because
their amnesia was much more severe than that of the other amnesic
patients and because both patients had sustained bilateral damage
to the amygdaloid complex, which has been linked to emotion and
emotional memory in humans and experimental animals (Cahill et
al., 1995; LeDoux, 1993).
Patient E. P. sustained extensive bilateral medial temporal lobe
damage, confirmed by MRI (Squire & Knowlton, 1995), which

included the hippocampus, perirhinal and parahippocampal cortices, the amygdaloid complex, and the inferotemporal gyrus. Aside
from his profound amnesia, E. P. also is anomic, scoring 42 of 60
on the Boston Naming Test (normal score > 50; Kaplan, Goodglass, & Weintraub, 1983). E. P. also exhibited some behavioral
evidence of frontal lobe dysfunction: On the Wisconsin Card
Sorting Test, 0 categories were sorted, and there were 51%
perseverative errors (Heaton, 1995); on the FAS Test of Word
Fluency, 18 words were produced, which fell below the 15th
percentile (Benton & Hamsher, 1976).
G. T. sustained extensive bilateral temporal lobe damage confirmed by MRI (Hamann et al., 1996). The damage extended

Table 2
Memory Test Performance
Patient

R.C.
N.F.
P.N.
J.W.
M.G.
H.W.
A.B.

PH.
L.J.

M
G.T.
E.P.

M
Control ( = 8)

Diagram
recall

3
4
2
4
0
6
4
3
3
3.2
0
0
0.0
20.6

Paired
associates

0
0
1
0
0
0
1
0
0
0.2
0
0
0.0
6.0

0
0
1
0
0
1

1
0
0
0.3
0
0
0.0
7.6

%Word
recall

3
2
1
2
2
0
2
1
0
1.4
0
0
0.0
8.9

19
36
29
28
33
31
33
27
40

%Word
recognition

50
words

50
faces

85
76
83
96
71
85
83
84
93

37
28
31
29
39
23
32
33
33

30
27
31
34
41
22
33
41
29

30.7

84.0

31.7

32.0

20
24

70
65

27
19

27
18

22.0

67.5

71

98

23.0
41.1

22.5
38.1

Note. The diagram recall score is based on delayed (12-min) reproduction of the Rey-Osterrieth
figure (Osterrieth, 1944; maximum score = 36). The average score for copying the figure was 28.8, a
normal score (Kritchevsky et al. 1988). The paired-associate scores are the number of word pairs
recalled on three successive trials (maximum score = 10/trial). The word recall score is the
percentage of 15 words identified correctly on five successive study-test trials (Rey, 1964). The word
recognition score is the percentage of 15 words identified correctly by yesno recognition across five
successive study-test trials. The scores for words and faces are based on 24-hr recognition tests of 50
words or 50 faces (modified from Warrington, 1984; maximum score = 50, chance = 25). The mean
scores for healthy controls and alcoholics shown for these tests are from Squire and Shimamura
(1986).

PERCEPTION AND MEMORY EM AMNESIA

through the anterior 7.0 cm of his left temporal lobe and the
anterior 5.0 cm of his right temporal lobe. The lesion included
bilaterally the hippocampus; entorhinal, perirhinal, and parahippocampal cortices; the amygdaloid complex; and the inferior, middle,
and superior temporal gyri. In addition to his profound amnesia,
G. T. is also anomic, scoring 18 of 60 on the Boston Naming Test
(normal score > 50; Kaplan et al., 1983).
Patients E. P. and G. T. were 72 and 59 years of age, respectively,
at the beginning of the study, and each had 12 years of education.
Immediate and delayed (12-min) recall of a short prose passage
averaged 1,5 and 0 segments, respectively (Gilbert et al., 1968;
maximum number of segments = 21). Scores on other memory
tests are shown in Tables 1 and 2. The mean score on the Dementia
Rating Scale was 112.0 (Mattis, 1976; maximum score = 144),
with most points being lost from the Memory subscale (mean
points lost = 16.0).
Control groups. The control and control delay groups were
employees or volunteers at the San Diego Veterans Affairs Medical
Center or were recruited from the retirement community of the
University of California, San Diego. The control group consisted of
6 men and 6 women who were selected to match the amnesic
patients with respect to the mean and range of their ages, years of
education, and scores on the Information and Vocabulary subtests
of the Wechsler Adult Intelligence ScaleRevised (WAIS-R;
Wechsler, 1981). They averaged 64.7 years of age (range = 5573), 13.6 years of education, and 20.4 and 53.0 on the Information
and Vocabulary subtests, respectively (amnesic patients = 18.1 and
51.4, respectively). Immediate and delayed recall of the short prose
passage averaged 6.7 and 5.0 segments, respectively.
An additional control group (control-delay) was tested after a
longer delay than that of the control group to bring their level of
memory performance into the performance range of the amnesic
patients. This group consisted of 6 men and 6 women selected to
match the amnesic patients with respect to the mean and range of
their ages, years of education, and scores on the Information and
Vocabulary subtests of the WAIS-R. They averaged 63.2 years of
age (range = 53-72), 14.6 years of education, and21.9 and 54.6 on
the Information and Vocabulary subtests, respectively (amnesic
patients = 18.1 and 51.4, respectively). Immediate and delayed
recall of the short prose passage averaged 7.1 and 6.3 segments,
respectively.

107

Materials and Procedure

The experiment consisted of a ratings phase, in which participants rated the emotionality of 40 color slides; a free-recall test, in
which participants attempted to recall the 40 slides they had seen;
and a yes-no recognition test in which 20 of the slides they had
seen were presented, together with 20 different distractor slides.
Finally, ratings of emotionality were collected for the 20 slides that
had served as distractor items in the recognition test.
Sixty color slides depicting various scenes and subject matter
were selected from the IAPS (Lang et al., 1988) to sample a wide
range of emotional content. The majority of these slides also had
been used as stimuli in the Bradley et al. (1992) study. The
procedure for the three groups was identical, except that the
amnesic and control groups were tested in a single session and the
control-delay group was tested with a 2-week delay between the
ratings phase and the other tests. After the 2-week delay, those in
the control-delay group were reminded that they had seen a slide
presentation in the previous session 2 weeks earlier. They were also
informed that the memory tests they would be taking would
concern the slides presented in that session.
Ratings phase. Slides were presented on a carousel slide
viewer. Participants were informed that they would be viewing a
series of pictures and that they were to rate each picture according
to the feeling they experienced while viewing it. No mention was
made of the memory tests that would follow.
Participants rated each of the 40 slides on two dimensions,
arousal (emotional intensity) and valence (degree of pleasant or
unpleasant affect), using the self-assessment mannequin (SAM)
rating system (Hodes, Cook, & Lang, 1985; Lang, 1980; see Figure
1). Arousal and valence each were assessed with the help of five
mannequin figures. Participants recorded their ratings by placing an
X on or between any of the five figures, which resulted in a possible
range of responses from 1 to 9. In the case of the arousal dimension,
the appearance of the corresponding SAM figures ranged from
having an active body and eyes wide open to having an inactive
body and closed eyes (see Figure 1). In the case of the valence
dimension, the appearance of the corresponding SAM figures
ranged from smiling with raised eyebrows to frowning with knitted
eyebrows (see Figure 1). Participants' ratings of emotion using
these two self-report scales have been demonstrated to covary

Figure 1. Scales used for arousal (top) and valence (bottom) ratings. For arousal, the scale assessed
the degree of emotional intensity. For valence, the scale assessed the degree of pleasant or unpleasant
affect.

108

HAMANN, CAHILL, AND SQUIRE

reliably with physiological reactions (Bradley et al., 1992; Greenwald, Cook, & Lang, 1989). Thus, arousal ratings covary reliably
with skin conductance responses, and valence ratings reliably
covary with facial electromyographic responses.
The two ratings scales were displayed on each page of a ratings
booklet (one page for each of 40 slides). For half the participants,
the upper scale assessed the arousal dimension and the lower scale
assessed the valence dimension; for the other half, the scales were
reversed. Because few ratings were placed between the mannequin
figures, the 9-point scale was transformed into a 5-point scale for
scoring, as described by Bradley et al. (1992).
Practice was given in the ratings procedure using two slides that
did not otherwise appear in the study (a slide of an umbrella and a
slide of a chocolate bar). Participants were informed that for each
slide they would have 12 s to make both ratings and that they
should attempt to record their initial emotional reaction to each
picture. It was emphasized further that they should try to use the
whole range of the ratings scale and that each rating should be
made on the basis of their true feelings.
To begin, the first practice slide was presented. After 12 s had
elapsed, the projector was advanced to a blank slide until the
participant was ready to rate the second practice slide. After the two
practice slides were rated, and additional instruction in the ratings
procedure was given if needed, the ratings phase proceeded in the
same manner as the practice phase. A card explaining the arousal
and valence ratings scales was always in view. In addition, the
amnesic patients were reminded periodically during the ratings
phase of the ratings procedure about the meaning of the ratings
scales.
For the ratings phase, the 40 slides were presented in a different
random order for each participant. Slides always were presented for
the full 12s, even when both ratings were completed quickly. In the
few cases in which the ratings were not completed within 12 s, the
projector was advanced to the blank slide and the participant
completed the ratings.
Recall test. When the ratings phase ended, participants were
tested for recall of the slides. They were instructed to write down in
any order a word or phrase describing each picture they remembered. They were told that they would be given 5 min to complete
this test.
Recognition test. After the free-recall test, a recognition test
was given in which 20 of the pictures presented in the ratings phase
were presented again as recognition targets together (in a pseudorandom order) with 20 new distractor pictures that matched the
targets -with respect to arousal, valence, and subject matter (using
the values for arousal and valence reported by Lang et al., 1988).
All 40 pictures in the recognition phase were digitized and
presented on a Macintosh Color Classic II computer. Each picture
measured 1 2 x 8 cm.
Participants were told that pictures would appear one at a time on
the computer screen, some of which had been presented in the
earlier ratings phase. If the picture had appeared in the ratings
phase, they were to press the button labeled yes (the z key for half
the participants and the / key for the other participants). Otherwise,
they were to press the button labeled no (the / key for half of the
participants and the z key for the other participants). Participants
were instructed to attempt to be as accurate as possible but to
respond as soon as they had decided because response times were
being recorded. The recognition test began approximately 7 min
after the end of the ratings phase. Each picture remained on the
screen until the participant pressed the yes or no key. Pressing
either key replaced the picture with a blank screen, which, after 2 s,
was followed by the next picture.
Final ratings phase. After the end of the recognition test, the
20 slides that had served as distractor iisms in the recognition test

were rated for arousal and valence, using the same procedure as in
the initial ratings phase.

Results
Arousal and Valence Ratings for the 60 Slides
The arousal and valence ratings for the 60 slides were
evaluated for the amnesic group, Patients E. P. and G. T., the
control group, the control-delay group, and a normative
sample of 87 undergraduates reported previously (Lang et
al., 1988). First, correlations were calculated between groups
for both arousal and valence ratings to determine whether
the groups gave the 60 slides the same pattern of ratings. All
correlations were significant at the .0001 level (df = 58).
Correlations were obtained by first calculating the mean
rating given each slide by all participants in each group and
then calculating for each pair of groups the correlation
between the mean ratings given the 60 slides. The Pearson
product-moment correlations for mean arousal ratings among
the amnesic, control, control-delay, and normative sample
groups ranged from .63 to .74, and the correlations for the
mean valence ratings ranged from .81 to .93. Patients E. P.
and G. T. gave arousal and valence ratings similar to the
other groups (the correlations for mean arousal ratings
between E; P. and G. T. and the other groups ranged from .35
to .49, ps < .01; for valence, the correlations ranged from
.78to.84,/is <.0001).
Although the groups assigned similar patterns of ratings
to the 60 slides, these groups (except for the control-delay
group) rated the 60 slides higher on the arousal dimension
than did the normative sample (see Tables 3 and 4), f(59) =
8.87, p < .0001, for the amnesic group; t(59) = 7.91, p <
.0001, for Patients E. P. and G. T.; and t(59) = 2.11, p < .04,
for the control group. These differences may be because the
participants in the current study were older than the undergraduates in the normative sample. In addition, the amnesic
group rated the 60 slides slightly higher on the arousal
dimension than did the control group or the control-delay
group (see Tables 3 and 4), ft(59) > 6.84, ps < .0001. For
the valence dimension, the amnesic group rated the 60 slides
similarly to the normative sample group, t(59) = 0.05, and
the control-delay group, f(59) = 0.84, but higher than the
control group, f(59) = 2.47, p < .02. Patients E. P. and G. T.
rated the 60 slides similarly to the ratings of the amnesic
group, ts(59) < 1.53, for arousal and valence.

Recall of 40 Slides
Free-recall responses were scored by comparing each
participant's written descriptions with the characteristics of
the 40 target recall slides. Two independent judges rated a
response correct if it could be linked clearly to a particular
slide that had been shown. Responses were not scored as
correct if they matched more than one slide (i.e., if the
response was not sufficiently specific to identify one particular slide). The majority of responses (87%) were scored as
correct or incorrect by both judges. Scoring conflicts between the two judges (13% of the responses) were resolved
by a third independent judge.

PERCEPTION AND MEMORY IN AMNESIA

109

Table 3
Arousal Ratings for 40 to-Be-Recalled Slides
Arousal quartile
Group
Control
Control-delay
Amnesic
Patients E.P. and G.T.

First
1.57
1.74
1.92
2.75

Second

0.18
0.29
0.19
0.45

2.70
2.72
3.00
3.50

0.21
0.31
0.24
0.50

Third
3.45
3.29
4.16
4.20

Fourth

0.22
0.30
0.20
0.20

4.42
4.14
4.83
4.80

0.15
0.25
0.11
0.20

Note. Numbers are means and the standard errors of the means. Arousal quartiles were constructed
by rank ordering slides according to each participant's arousal ratings. Each quartile contained 10
slides.
The recall results were analyzed according to the arousal
score given each slide. The 40 target recall slides were rank
ordered for each participant according to each participant's
arousal ratings. (The ranks for slides given identical arousal
ratings by the participant were decided on the basis of the
arousal means for the participant's group.) Figure 2 shows
the results for the amnesic, control, and control-delay
groups. Neither E. P. nor G. T. was able to recall any studied
slides. Table 3 shows the mean arousal ratings that corresponded to each arousal quartile in Figure 2.
A mixed-factorial analysis of variance (ANOVA) was first
conducted with group (amnesic vs. control) as the betweensubjects variable and arousal quartile as the within-subjects
variable. The proportion of correctly recalled slides was the
dependent variable. There was an effect of group, F(l, 19) =
64.29, MSB = 0.022, p < .0001, and of arousal, F(3, 57) =
8.10, MSE = 0.012, p < .0001, but no interaction between
group and arousal, F(3, 57) < 1. Thus, the amnesic group
was impaired relative to the control group, and both groups
showed a similar beneficial effect of arousal on recall.
Next, the same analysis was conducted between the
amnesic group and the control-delay group. There was an
effect of arousal, F(3, 57) = 7.09, MSE = 0.010, p < .001,
but no effect of group, F(l, 19) = 1.52, MSE = 0.015, p >
.23, and no interaction between group and arousal, F(3,
57) < 1. The failure to find a Group X Arousal interaction
argues against the possibility that the absence of the same
interaction in the amnesic versus control group comparison
was attributable to the markedly different levels of recall.
Next, the effect of arousal on recall was examined
separately in each group across arousal quartiles. For the
amnesic group, the effect of arousal on recall was not
significant, F(3, 24) = 1.84, MSE = 0.013, p = .17. A test
for the presence of a linear trend across arousal quartiles for

recall in the amnesic group fell short of significance

(p < .09). The effect of arousal on recall was significant for
both the control group, F(3, 33) = 8.03, MSE = 0.012, p <
.001, and the control-delay group, F(3, 33) = 7.05, MSE =
0.009, p<. 001.
Recognition of 20 Slides
The recognition data were analyzed according to whether
a slide belonged to the low- or high-arousal category. The 20
target and 20 distractor recognition test slides were rank
ordered for each participant according to each participant's
arousal ratings. (The ranks for slides given identical arousal
ratings were decided on the basis of the arousal means for
the participant's group.) The 10 slides in each set (target or
distractor) given the highest ratings were designated as
high-arousal slides; the other 10 were designated as lowarousal slides. The recognition data were partitioned into
two arousal categories (low and high arousal) rather than
into quartiles (as with the recall data), so that the same
number (10) of slides were in each arousal category for both
the recall and recognition data.
Figure 3 shows die proportion of recognition hits and
false alarms (FAs) for the amnesic, control, and controldelay groups as a function of arousal score. Table 4 shows
the mean arousal ratings dial corresponded to the low- and
high-arousal categories in Figure 3.
Three mixed-factorial ANOVAs were conducted with
group (amnesic vs. control) as the between-subjects variable, arousal (low vs. high) as the within-subjects variable,
and proportion of hits, FAs, and corrected recognition scores
as the dependent measures, respectively. The amnesic group
was impaired relative to the control group on all three
measures. For all three measures, there was an effect of

Table 4
Arousal Ratings for 20 Target Slides and 20 Distractor Slides in the Recognition Test
Target
Group
Control
Control-delay
Amnesic
Patients E.P. and G.T.

Distractor

Low arousal

High arousal

Low arousal

High arousal

2.02
2.42
2.51
3.00

3.96
3.87
4.69
4.65

2.37
2.48
2.88
2.80

3.98
3.84
4.72
4.65

0.17
0.31
0.19
0.40

0.15
0.27
0.12
0.15

0.17
0.24
0.17
0.30

0.16
0.24
0.11
0.25

Note. Numbers are means and the standard errors of the means. The low- and high-arousal
categories were constructed by rank ordering slides according to each participant's arousal ratings.
Each arousal category contained 10 slides.

110

HAMANN, CAHILL, AND SQUIRE

AROUSAL QUARTILE

Figure 2. Recall scores for 40 slides obtained by the control

(CON; n = 12), amnesic (AMN; n = 9), and control-delay (CONDELAY; n = 12) groups as a function of the arousal score given
each slide (first, second, third, or fourth quartile). Error bars
indicate the standard error of the mean.

group, Fs(l, 19) - 5-55-12.09, ps < .05; an effect of

arousal, Fs(l, 19) = 5.39-12.37, ps < .05; and no Group X
Arousal interaction, Fs(l, 19) = 0.64-2.11, ps > .16.
Next, the same analysis was conducted between the
amnesic group and the control-delay group. Memory performance was similar in both groups. There was no effect of
group for hits, FAs, or corrected recognition scores,
F(l, 19) < 1.60. The effect of arousal was significant for hits
and corrected recognition scores, F(l, 19) > 8.3, and there
was no Group X Arousal interaction (Fs < 1.02). The
absence of a Group X Arousal interaction argues against the

possibility that the absence of the same interaction in the

amnesic versus control group comparison was attributable to
differing levels of recognition performance or to ceiling
effects in the control group.
After these group comparisons, the effect of low versus
high arousal on hits, FAs, and corrected recognition was
examined separately in each group. For the amnesic group,
there was a marginal effect of arousal on hits, f(8) = 1.94,
p < .09, no effect on FAs, /(8) = \M,p > .14, and an effect
on corrected recognition scores, f(8) = 2.64, p < .05. For the
control group, there was a marginal effect of arousal on hits,
r(ll) = 2.16,/j < .054, no effect on FAs, f(ll) = l.6Q,p>
.14, and a marginal effect on corrected recognition scores,
t(ll) = 2.06, p < .06. For the control-delay group, there was
an effect of arousal on hits, t(ll) = 2.20, p < .05, no effect
on FAs (p > .1), and an effect on corrected recognition
scores, t(U) = 336,p < .01.
Recognition performance for Patients E. P. and G. T. was
near floor, at an overall level only marginally above chance
(hit rates and FAs were converted to d'): mean d' = .54,
t(l) = 3.92 against 0, p < .08 (one-tailed test). The mean hit
rate was .40 .00 and .55 .05 for the low- and
high-arousal slides, respectively; the corresponding FA rates
were . 10 .00 and .45 .05, respectively. In contrast to the
amnesic, control, and control-delay groups, who showed a
memory benefit from arousal, the corrected recognition
scores of Patients E. P. and G. T. were (nonsignificantly)
lower for high-arousal slides than for low-arousal slides
(.10 .10 and .30 .00, respectively).
Effects of Valence
Valence had virtually no effect on memory, independent
of arousal scores. This result is understandable given the

LOW AROUSAL
HIGH AROUSAL

DC

CON

AMN CON-DELAY

HITS

CON

AMN CON-DELAY

FALSE ALARMS

Figure 3. Recognition hits and false alarms (FAs) for the control (CON; n 12), amnesic (AMN;
n = 9), and control-delay (CON-DELAY; n = 12) groups for low-arousal (open bars) and
high-arousal (dark bars) slides. Error bars indicate the standard error of the mean. Corrected
recognition scores (hits minus FAs) for the control group were .83 .05 and .92 .04 for the lowand high-arousal slides, respectively; the corresponding values for the amnesic group were .47 .10
and .66 .09, respectively, and the corresponding values for the control-delay group were .38 .06
and .50 .05, respectively.

111

PERCEPTION AND MEMORY IN AMNESIA

Table 5
Reaction Times for the 20 Target Slides and 20 Distractor Slides in the Recognition Test
Targets
Group
Control
Control-delay
Amnesic
Patients E.P. and G.T.

Low arousal
1,119
1,427
1,360
3,282

206
110
149
1,277

Distractors
High arousal
963
1,663
1,292
2,256

84
145
122
176

Low arousal
1,052
1,843
1,435
3,101

86
115
143
1,888

High arousal
1,178
2,269
1,722
2,739

100
178
192
1,511

Note. Numbers are means and the standard errors of the means. The low- and high-arousal
categories were constructed by rank ordering slides according to each participant's arousal ratings.
Each arousal category contained 10 slides. Only reaction times for correct responses (hits and false
alarms) were analyzed.

marked curvilinear (quadratic) relationship between valence

and arousal ratings in both the current study (quadratic r
values between valence and arousal ratings for the 60 slides
ranged from .53 to .74, p < .0001, for the amnesic, control,
and control-delay groups and for Patients E. P. and G. T.) and
in previous studies (Bradley et al., 1992; Lang et al., 1988).
Specifically, both highly unpleasant (low-valence) and highly
pleasant (high-valence) slides were rated as being more
arousing than neutral slides.
As a result, when the slides in each arousal category (four
arousal categories in the case of the recall test and two in the
case of recognition) were partitioned for each participant
into equal numbers of high- and low-valence slides, the
arousal ratings for the high- and low-valence slides were the
same. Moreover, when valence was included as a variable in
the ANOVAs that had been carried out to analyze the recall
and recognition test results, no main effects of valence were
obtained. In addition, there were no interactions involving
valence and the group variable.
Reaction Time Analysis
Only correct trials (hits and correct rejections) were used
to analyze the recognition reaction time data. For each
group, a within-subjects ANOVA was conducted with slide
status (target vs. distractor), arousal, and valence as variables. In addition, reaction times greater than 5 s were
omitted from analysis (3% of responses). Table 5 shows
mean reaction times for each group as a function of slide
status and arousal. The reaction time measure, in contrast to
accuracy, was relatively insensitive to arousal. The amnesic
group, F(l, 8) = 5.63, MSE = 203, 721, p < .05, and the
control-delay group, F(l, 11) = 14.79, MSE = 423, 417,
p < .01, both responded faster to old (target) slides than to
new (distractor) slides (reflecting a savings in encoding time
for old slides; 1,326 95 vs. 1,578 121 and 1,545 92
vs. 2,056 109 for the amnesic and control-delay groups,
respectively). The control group also responded (numerically)fastertooldslides,butnotsignificantlyso(l,041 111
vs. 1,115 66, F < 1). There were no other main effects or
interactions, with the single exception of a main effect of
arousal in the control-delay group, F(l, 11) = 17.61, MSE =
149, 515, p < .01. High-arousal slides (both targets and
distractors) were responded to more slowly than low-arousal
slides (1,634 85 and 1,966 122, respectively).

Comparison of Patients With and Without

Korsakoff's Syndrome
The performance of the 5 patients with Korsakoff's
syndrome on emotional ratings of the 60 slides and on the
recall and recognition tests was compared with the performance of the 4 non-Korsakoff amnesic patients. The two
groups rated the 60 slides similarly overall for both arousal
(3.53 .11 vs. 3.66 .08 for the Korsakoff and nonKorsakoff patients, respectively), r(59) = 1.60, p > .11, and
for valence (3.16 .16vs.3.15 .15 for the Korsakoff and
non-Korsakoff patients, respectively), r(59) < 1. Furthermore, the pattern of ratings across the 60 slides was highly
similar between the two groups (rs = .71 for between
groups on arousal ratings, p < .0001, and .83 for valence
ratings, p< . 0001).
To compare memory performance between the two groups,
we conducted four mixed-factorial ANOVAs with proportion correct recall, recognition hits, FAs, and corrected
recognition score as the dependent variables. The betweensubjects variable for each ANOVA was subject group, and
the within-subjects variables were arousal and valence. The
Korsakoff amnesic patients performed similarly to the
non-Korsakoff patients on each measure, with no effects of
group or interactions involving group and the arousal or
valence variables. The exception was a three-way interaction of group, arousal, and valence for recognition hits, F(1,
7) = 28.64, MSE = 0.004, p < .01. This interaction reflected
the fact that the amnesic patients with Korsakoff s syndrome
in general had a slightly lower proportion of hits than the
non-Korsakoff patients, except for highly arousing, pleasant
slides. For these slides, the patients with Korsakoff's syndrome outperformed the non-Korsakoff patients (.84 .08
vs. .65 .17 correct, respectively).

General Discussion
We examined whether perception of emotional stimuli is
normal in amnesia and whether emotional arousal has the
same enhancing effect on memory in amnesic patients as it
has in healthy controls. With respect to perception of
emotional stimuli, the amnesic group, as well as amnesic
Patients E. P. and G. T., rated the 60 slides similarly to our
controls and the normative sample originally reported by
Lang et al. (1988). The amnesic group and Patients E. P. and

112

HAMANN, CAHILL, AND SQUIRE

G. T. did tend to give higher arousal and valence ratings to

the slides than did controls and the normative sample group,
but these differences were small (see Tables 3 and 4) and the
overall pattern of slide ratings was remarkably similar across
all groups. The finding that Patients E. P. and G. T., who
have bilateral damage to the amygdala, gave normal emotionality ratings is consistent with the previous finding that
Patient B. P., a patient with bilateral amygdala damage
resulting from Urbach-Wiethe disease, was unimpaired in
rating the arousal associated with an emotional story (Cahill
etal, 1995).
Arousal improved recall and recognition performance in
all groups except Patients E. P. and G. T., who exhibited little
or no detectable memory for the slides. Although the
amnesic patients were severely impaired on both recall and
recognition compared with the control group, their memory
was enhanced by high arousal in proportion to the enhancement exhibited by the control group. In addition, when the
amnesic group was compared with the control-delay group,
which had been matched to the performance of the amnesic
group by testing after a long delay, the two groups exhibited
the same memory enhancement as a function of arousal.
Indeed, for recognition, the amnesic group exhibited numerically more benefit from arousal than the control-delay
group. For recall, the groups also benefited similarly from
arousal, and there was no interaction with arousal in any
group comparison. For the recall measure, the test for a
linear trend for an arousal effect did not reach significance
when the amnesic group was examined by itself (p < .09),
likely because of floor effects in the amnesic group. Our
results for free recall are consistent with previous findings
that patients with left unilateral temporal lobectomy exhibit
the same benefit in free recall for emotional versus neutral
words as controls, despite an overall recall deficit for the
patients (Phelps, Ziv, & Labar, 1995). Thus, the memory
enhancement effect associated with arousal appears to be
intact in amnesia, and this effect extends to both recall and
recognition memory.
The lack of a memory enhancement effect in Patients E. P.
and G. T. cannot easily be interpreted. Both E. P. and G. T.
have bilateral damage to the amygdala, and when bilateral
amygdala damage is present emotional arousal may not
enhance memory (Cahill et al., 1995). However, the recognition performance of these patients was also near floor, which
would have made it difficult to detect an enhancement effect.
Either or both of these variables could account for the lack of
a memory enhancement effect in these 2 patients.
The comparison between recall and recognition performance in the amnesic and control-delay groups also is
notable. To our knowledge, this is the first demonstration
that free recall and recognition of pictorial information are
impaired in amnesia proportional to their difficulty for
controls (see Figures 2 and 3). Previous work (Haist.
Shimamura, & Squire, 1992) established that free recall and
recognition are impaired equivalently in amnesia when word
lists were used. The current finding provides further evidence that recognition in amnesia is not being supported by
memory systems (such as priming) that are intact in amnesia
(also see Squire & Knowlton, 1995). The deficit in recogni-

tion was about what it should have been given the deficit in
free recall.
The effect of valence was minor in the current study once
the effect of arousal on memory had been accounted for.
This result is understandable, given that valence and arousal
ratings for the 60 slides were correlated highly. The effect of
valence was limited to slightly higher corrected recognition
performance for low-valence items in the control and
control-delay groups (there was no effect in the amnesic
group), and an interaction between arousal and valence for
FAs in the amnesic versus control group comparison. The
minor effect of valence on recall and recognition in the
current study accords well with similar findings by Bradley
et al. (1992) for recall and recognition in college students.
Reaction time to recognition items was relatively insensitive to the effects of emotional arousal. Bradley et al. (1992)
did report an effect of emotional arousal on reaction time in
picture recognition. Specifically, hit responses to higharousal slides were faster than hit responses to low-arousal
slides, and correct rejections were slower for high-arousal
slides than for low-arousal slides. Perhaps in the current
study participants emphasized accuracy over speed.
The results of the current study suggest that emotional
judgments are intact in amnesia, even in patients with
bilateral damage to the amygdala. Previous clinical and
anecdotal reports have suggested that patients with Korsakoffs syndrome often are abnormal in their emotional
responses (Rapaport, 1961; Talland, 1965). However, our 5
patients with Korsakoff s syndrome rated the emotional
arousal and valence of slides highly similarly to the nonKorsakoff patients, and the beneficial effect of emotional
arousal on memory was similar in both groups. As considered previously (Shimamura, Jernigan, & Squire, 1988),
alcoholic Korasakoff s syndrome is variable with respect to
the severity of memory impairment, frontal lobe signs, and
other neuropsychological signs of cortical dysfunction. Our
patients were selected because they had a memory impairment disproportionate to any other intellectual impairment.
In summary, amnesic patients experienced the same
memory enhancement from emotional arousal as healthy
controls. The results suggest that the emotional arousal
effect is modulated by brain structures that are intact in
amnesia. The amygdala, which is spared in all the amnesic
patients except Patients G. T. and E. P., is a prime candidate
for the source of the emotional arousal effect, based on
evidence from both animals and humans (Davidson &
Sutton, 1995; McGaugh, 1989).

References
Benton, A. L., & Hamsher, K. deS. (1976). Multilingual Aphasia
Examination. Iowa City: University of Iowa Press.
Bradley, M. M., Grcenwald, M. K., Petty, M. C, & Lang, P. J.
(1992). Remembering pictures: Pleasure and arousal in memory.
Journal of Experimental Psychology: Learning, Memory, and
Cognition, 18, 379-390.
Cahill, L., Babinsky, R., Markowitsch, H. J., & McGaugh, J. L.
(1995). The amygdala and emotional memory. Nature, 377,
295-296.

PERCEPTION AND MEMORY IN AMNESIA

Cahill, L., Prins, B., Weber, M., & McGaugh, J. L. (1994).
f3-Adrenergic activation and memory for emotional events.
Nature, 371, 702-704.
Christiansen, S.-A., & Loftus, E. F. (1987). Memory for traumatic
events. Applied Cognitive Psychology, 1, 225-239.
Craik, F. I. M., & Blankstein, K. R. (1975). Psychophysiology and
human memory. In P. H. Venables & M. J. Christie (Eds.),
Research in psychophysiology (pp. 388-417). New York: Wiley.
Davidson, R. J., & Sutton, S. K. (1995). Affective neuroscience:
The emergence of a discipline. Current Opinion in Neurobiology, 5, 217-224.
Davis, M. (1994). The role of the amygdala in emotional learning.
International Review of Neurobiology, 36, 225266.
Douglas, J. J., & Wilkinson, D. A. (1993). Evidence of normal
emotional responsiveness in alcoholic Korsakoff s syndrome in
the presence of profound memory impairment. Addiction, 88,
1637-1645.
Gilbert, J., Levee, R., & Catalano, K. (1968). A preliminary report
on a new memory scale. Perceptual and Motor Skills, 27,
277-278.
Greenwald, M. K., Cook, E. W., & Lang, P. J. (1989). Affective
judgment and psychophysical response: Dimensional covariation in the evaluation of pictorial stimuli. Journal of Psychophysiology, 3, 51-64.
Haist, F., Shimamura, A. P., & Squire, L. R. (1992). On the
relationship between recall and recognition memory. Journal of
Experimental Psychology: Learning, Memory, and Cognition,
18, 691-702.
Hamann, S. B., Stefanacci, L., Squire, L. R., Adolphs, R., Tranel,
D., Damasio, H., & Damasio, A. (1996). Recognizing facial
emotion. Nature, 379, 497.
Heaton, R. K. (1995). Wisconsin Card Sorting Test manual.
Odessa, FL: Psychology Assessment Resources.
Heuer, F., & Reisberg, D. (1990). Vivid memories of emotional
events: The accuracy of remembered minutiae. Memory &
Cognition, 18, 496-506.
Hodes, R. L., Cook, E. W., & Lang, P. J. (1985). Individual
differences in autonomic response: Conditioned association or
conditioned fear? Psychophysiology, 22, 545-560.
Janowsky, J. S., Shimamura, A. P., & Squire, L. R. (1989). Source
memory: Impairment in patients with frontal lobe lesions.
Neuropsychologia, 27, 1043-1056.
Kaplan, E. F., Goodglass, H., & Weintraub, S. (1983). The Boston
Naming Test. Philadelphia: Lea & Febiger.
Kritchevsky, M., Squire, L. R., & Zouzounis, J. A. (1988).
Transient global amnesia: Characterization of anterograde and
retrograde amnesia. Neurology, 38, 213-219.
Lang, P. J. (1980). Behavioral treatment and bio-behavioral assessment: Computer applications. In J. B. Sidowski, J. H. Johnson, &
T. A. Williams (Eds.), Technology in mental health delivery
systems (pp. 119-137). Norwood, NJ: Ablex.
Lang, P. J., Bradley, M. M., & Cuthbert, B. N. (1990). Emotion,
attention, and the startle reflex. Psychological Review, 97,
377-398.
Lang, P. J., Ohman, A., & Vaitl, D. (1988). The International
Affective Picture System [Photographic slides]. Gainesville:
University of Florida, Center for Research in Psychophysiology.
LeDoux, J. E. (1993). Emotional memory systems in the brain.
Behavioural Brain Research, 58, 69-79.
Markowitsch, H., Kessler, J., & Denzler, P. (1986). Recognition
memory and psychophysiological responses to stimuli with

113

neutral or emotional content: A study of Korsakoff patients and

recently detoxified and longterm abstinent alcoholics. International Journal of Neuroscience, 29, 135.
Matlin, M., & Stang, D. (1978). The Pollyannaprinciple: Selectivity in language, memory, and thought. Cambridge, MA: Schenkman.
Mattis, S. (1976). Dementia Rating Scale. In R. Bellack & B.
Keraso (Eds.), Geriatric psychiatry (pp. 77-121). New York:
Grune & Stratton.
McGaugh, J. L. (1989). Involvement of hormonal and neuromodulatory systems in the regulation of memory storage. Annual
Review of Neuroscience, 12, 255-287.
McGaugh, J. L., Introini-Collison, I. B., Cahill, L. F., Castellano,
C., Dalmaz, C., Parent, M. B., & Williams, C. L. (1993).
Neuromodulatory systems and memory storage: Role of the
amygdala. Behavioural Brain Research, 58, 81-90.
Oscar-Berman, M., Hancock, M., Mildworf, M. S., Hutner, N., &
Weber, D. A. (1990). Emotional perception and memory in
alcoholism and aging. Alcoholism: Clinical and Experimental
Research, 14, 383-393.
Osterrieth, P. A. (1944). Le test de copie d'une figure complexe
[The test of copying a complex figure]. Archives de Psychologic,
30, 206-356.
Phelps, E. A., Ziv, O., & Labar, K. S. (1995). Recall and affect
following unilateral temporal lobectomy. Society for Neuroscience Abstracts, 21, 755.
Rapaport, D. (1961). Emotions and memory. New York: Science
Editions.
Rey, A. (1964). L'examen clinique psychologic [The clinical exam
in psychology]. Paris: Presses Universitaires de France.
Russell, J. (1980). A circumplex model of affect. Journal of
Personality and Social Psychology, 39, 1161-1178.
Shimamura, A. P., Jernigan, T. L., & Squire, L. R. (1988).
Korsakoff s syndrome: Radiological (CT) findings and neuropsychological correlates. Journal of Neuroscience, 8, 4400-4410.
Squire, L. R., Amaral, D. G., & Press, G. A. (1990). Magnetic
resonance measurements of hippocampal formation and mammillary nuclei distinguish medial temporal lobe and diencephalic
amnesia. Journal of Neuroscience, 10, 3106-3117.
Squire, L. R., & Knowlton, B. J. (1995). Learning about categories
in the absence of memory. Proceedings of the National Academy
of Sciences, 92, 12470-12474.
Squire, L. R., & Shimamura, A. P. (1986). Characterizing amnesic
patients for neurobehavioral study. Behavioral Neuroscience,
100, 866-877.
Squire, L. R., & Zola-Morgan, S. (1991). The medial temporal lobe
memory system. Science, 253, 1380-1386.
Stuss, D. T., Gow, C. A., & Hetherington, C. R. (1992). "No longer
Gage": Frontal lobe dysfunction and emotional changes. Journal of Consulting and Clinical Psychology, 60, 549559.
Talland, G. A. (1967). Deranged memory. San Diego, CA: Academic Press.
Warrington, E. K. (1984). Recognition Memory Test. Windsor,
England: FER-Nelson.
Wechsler, D. (1981). The Wechsler Mult Intelligence ScaleRevised. New York: Psychological Corporation.

Received March 1, 1996

Revision received May 28, 1996
Accepted May 31,1996