Neuropsychologia 45 (2007) 1305–1317

Role of the amygdala in decisions under ambiguity and decisions

under risk: Evidence from patients with Urbach-Wiethe disease
Matthias Brand a,∗ , Fabian Grabenhorst a , Katrin Starcke a ,
Marie M.P. Vandekerckhove a,b,c , Hans J. Markowitsch a
a Department of Physiological Psychology, University of Bielefeld, Bielefeld, Germany
b Department of Psychology, University of Antwerpen, Antwerpen, Belgium
c Department of Medicine, University of Antwerpen, Antwerpen, Belgium

Received 30 January 2006; received in revised form 22 September 2006; accepted 27 September 2006
Available online 27 October 2006

Abstract
Various neuropsychological studies have shown that decision-making deficits can occur in a wide range of patients with brain damage or
dysfunctions. Decisions under ambiguity, as measured with the Iowa Gambling Task, primarily depend on the integrity of the ventromedial
prefrontal cortex and the amygdala, as well as on further brain regions such as the somatosensory cortex. However, little is known about the specific
role of these structures in decisions under risk measured with tasks that offer explicit rules for gains and losses and winning probabilities, for example,
the Game of Dice Task. We aimed to investigate the potential role of the amygdala for decisions under risk. For this purpose, we examined three
patients with Urbach-Wiethe disease—a rare syndrome associated with selective bilateral mineralisation of the amygdalae. Neuropsychological
performance was assessed with the Iowa Gambling Task (decisions under ambiguity), the Game of Dice Task (decisions under risk), and an
extensive neuropsychological test battery focussing on executive functions. Furthermore, previous studies found relationships between generating
skin conductance responses and deciding advantageously in the Iowa Gambling Task. Accordingly, we recorded skin conductance responses during
both decision tasks as a measure of emotional reactivity. Results indicate that patients with selective amygdala damage have lower scores in both
decisions under ambiguity and decisions under risk. Decisions under risk are especially compromised in patients who also demonstrate deficits in
executive functioning. In both gambling tasks, patients showed reduced skin conductance responses compared to healthy comparison subjects. The
results suggest that deciding advantageously under risk conditions involves both the use of feedback from previous trials, as required by decisions
under ambiguity, and in addition, executive functions.
© 2006 Elsevier Ltd. All rights reserved.

Keywords: Decision-making; Iowa Gambling Task; Game of Dice Task; Emotional processing; Prefrontal cortex

1. Introduction clear/obvious probabilities. These types of decisions – at least

Deciding between different options is a very important func-
tion in everyday life and disturbances of decision-making can
result in severe social, financial and health problems. In real
life, decision situations differ in their degree and probability
of associated reward and punishment. In decisions under ambi-
guity individuals have to decide between different options, but
the outcome of their choices is uncertain and not defined by
∗ Corresponding author at: Department of Physiological Psychology, Univer-
sity of Bielefeld, P.O. Box 100131, 33501 Bielefeld, Germany.
Tel.: +49 521 106 4488; fax: +49 521 106 6049.
E-mail address: m.brand@uni-bielefeld.de (M. Brand).
in patients with brain lesions – primarily depend on the integrity
of the ventromedial prefrontal cortex and the amygdala as well as
on structures involved in limbic circuits (e.g., Bechara, Damasio,
& Damasio, 2000, 2003; Bechara & Van Der Linden, 2005).
According to the somatic marker hypothesis (e.g., Damasio,
1996), subjects have to follow their own feelings and hunches
for optimal decision-making (Bechara, 2001). Although dis-
cussed controversially (see Maia & McClelland, 2004, 2005),
in the process of decision-making, the amygdala is suggested to
be a critical structure as being involved in processing primary
inducers and emotional arousal associated with anticipation
of rewards and punishments (e.g., Kringelbach, 2005; Phelps,
2006; Phelps & LeDoux, 2005). Decisions under ambiguity are
often tested using the Iowa Gambling Task (abbreviated with

0028-3932/$ – see front matter © 2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.neuropsychologia.2006.09.021
1306 M. Brand et al. / Neuropsychologia 45 (2007) 1305–1317
“gambling task” in the following) (Bechara, Damasio, Damasio,
& Anderson, 1994; Bechara, Tranel, & Damasio, 2000). In this
task, subjects are asked to select a card from one of four card
decks. After card selection, a fictitious gain is displayed, irregu-
larly accompanied by a loss. Two of the four decks are coupled
with high gains but even higher losses. Choosing from these
disadvantageous decks will lead to a negative net-balance in
the long run. The other two decks will result in small gains
but even smaller losses (advantageous decks). Thus, choosing
from these advantageous decks will lead to a positive overall
balance. Although participants are told that some card decks are
“better” than others, they are not explicitly informed about the
reward/punishment contingencies. Thus, the possible choices
are full of ambiguity and participants have to learn to avoid the
disadvantageous card decks using the feedback from previous
trials.
Using this task, decision-making deficits have been revealed
in a wide range of neurological patients suffering from frontal
lobe damage (especially when lesion foci centred around the
orbitofrontal/ventromedial section of the prefrontal cortex; e.g.,
Bechara, Tranel, et al., 2000). Furthermore, psychological
patient populations were found to be deficient relative to healthy
individuals in gambling task performance, such as patients
suffering from substance dependencies (e.g., Bechara, 2005;
Bechara & Damasio, 2002; Bechara et al., 2001; Bechara, Dolan,
& Hindes, 2002; Bolla et al., 2003; Bolla, Eldreth, Matochik, &
Cadet, 2005; Fishbein et al., 2005; Verdejo, Aguilar de Arcos, &
Perez Garcia, 2004; Whitlow et al., 2004), or patients with obses-
sive compulsive disorder, schizophrenia, pathological gambling,
anorexia nervosa, suicide attempters, and other patients with
neuropsychiatric symptoms (e.g., Bark, Dieckmann, Bogerts,
& Northoff, 2005; Cavedini et al., 2004; Cavedini, Riboldi,
D’Annucci et al., 2002; Cavedini, Riboldi, Keller, D’Annucci,
& Bellodi, 2002; Goudriaan, Oosterlaan, de Beurs, & van den
Brink, 2005; Jollant et al., 2005) (an excellent review of studies
with the gambling task is found in the article by Dunn, Dalgleish,
& Lawrence, 2006).
Knowledge about the role of the amygdala in decisions
under ambiguity comes from studies which examined patients
with selective damage of the amygdalae with the gambling
task. For instance, Bechara, Damasio, Damasio, and Lee (1999)
studied gambling task performance of five patients suffering
from Urbach-Wiethe disease (UWD) (n = 1), encephalitis dur-
ing childhood (n = 2) or herpes simplex encephalitis during
adulthood (n = 2), all having lesions comprising bilateral medial
temporal lobes and being more or less restricted to the amyg-
dalae. The authors compared gambling task performance of
these patients with performance of healthy subjects as well as
with that of patients with selective damage to the ventrome-
dial section of the prefrontal cortex. The behavioural data of
both patient groups appeared very similar indicating poor gam-
bling task performance compared to the healthy participants.
The authors also recorded skin conductance responses (SCRs),
as a measure of emotional reactivity, while the subjects per-
formed the task. Both, amygdala-damaged patients and patients
with ventromedial lesions had significantly lower anticipatory
SCRs (that is SCR during the time period before a choice) than
the normal comparison subjects. SCRs following each choice
differed between patient groups: Amygdala-damaged patients
generated significantly lower SCRs after both types of choices,
selections of decks that led to reward as well as after selec-
tions of cards that were followed by punishments, whereas
ventromedial PFC patients showed similar responses compared
to healthy individuals. Four of five patients with ventromedial
lesions generated SCRs within the range of the healthy com-
parison subjects’ scores. The authors concluded that the lack of
SCR generation after choice in amygdala-damaged patients may
reflect that gaining or losing money in the gambling task does
not evoke a somatic state in individuals with amygdala dam-
age. This, as the authors stated, is in accordance with real life
decision-making deficits of those patients (as described in Tranel
& Hyman, 1990). SCRs are seen as reflecting general arousal
associated with processing emotional stimuli (Boucsein, 1992;
Venables & Christie, 1980). Recording SCRs during gambling
task performance was done by a few studies and most of them
replicated the finding of the studies by Bechara and colleagues
(e.g., Bechara & Damasio, 2002; Bechara et al., 2002; Bechara,
Tranel, Damasio, & Damasio, 1996) who revealed higher antic-
ipatory SCRs before choosing a disadvantageous option relative
to those before choosing an advantageous card deck in healthy
subjects (e.g., Crone, Somsen, Van Beek, & Van Der Molen,
2004; Hinson, Jameson, & Whitney, 2002). The results were
interpreted as the higher anticipatory SCRs preceding disad-
vantageous decisions act as warning signals from the periphery
that gradually lead to avoiding the disadvantageous alternatives
and preferring the advantageous options. Other studies found
that generating SCRs after the feedback is delivered (potentially
indicating appraisal of the decision) may be more important to
perform well in the gambling task than the anticipatory SCRs.
For example, Suzuki, Hirota, Takasawa, and Shigemasu (2003)
found that feedback SCRs were higher after choosing a disad-
vantageous option than an advantageous and were higher for
punishments than for rewards. However, it is still a topic of
debate whether or not SCRs, as measures of emotional arousal,
guide future decisions and whether anticipatory and/or feedback
SCRs are primarily important for successful gambling task per-
formance (see the critical review by Dunn et al., 2006).
In contrast to the decisions measured in the gambling task,
many decisions in real life can be made on the basis of probabil-
ities and explicit knowledge about options and their associated
rewards and punishments (e.g., the amount of gains and losses).
Such decisions are termed “decisions under risk” (e.g., Bechara,
2004; Brand, Labudda, & Markowitsch, 2006). We have recently
developed a gambling task, the Game of Dice Task (abbre-
viated with “dice task” in the following) that is believed to
simulate those decisions under risk. Here, subjects are asked
to predict the outcome of a dice throw (see detailed descrip-
tion of the task in Section 2). This task offers explicit rules for
gains and losses and obvious winning probabilities. Thus, par-
ticipants are allowed the possibility of utilising explicit clues to
optimise their performance (e.g., by calculating the risk asso-
ciated with each option). In a series of studies with patients
suffering from neurological diseases (e.g., Morbus Parkinson,
Korsakoff’s syndrome) or psychological disorders (e.g., patho-
 M. Brand et al. / Neuropsychologia 45 (2007) 1305–1317
logical gambling), we found that performance in decisions under
risk was associated with both specific executive functions (e.g.,
categorisation, set-shifting) and processing feedback from pre-
vious trials within the task (Brand, Fujiwara et al., 2005; Brand,
Kalbe et al., 2005; Brand, Labudda et al., 2004). In accor-
dance with these behavioural studies and results obtained from
neuroimaging investigations (recent review in Krain, Wilson,
Arbuckle, Castellanos, & Milham, 2006), we suggested that
decisions under risk, as measured by the dice task, rely on intact
fronto-striatal loops connecting mesencephalic dopaminergic
structures (substantia nigra, ventral tegmental area) with both the
dorsal striatum and the dorsolateral section of the prefrontal cor-
tex (the so-called cognitive loop; Alexander & Crutcher, 1990;
Alexander, Crutcher, & DeLong, 1990) as well as limbic struc-
tures such as the amygdala, the nucleus accumbens as well as the
ventromedial and the orbitofrontal part of the prefrontal cortex
(the so-called limbic loop according to Alexander & Crutcher,
1990; Alexander et al., 1990). The relevance of the dorsolateral
prefrontal cortex for dice task performance is indirectly con-
firmed by a case study of a female patient who suffered from
severe decision-making deficits in real life following a foramen
of Monro cyst removal (Brand, Kalbe et al., 2004). She was
severely impaired relative to control subjects in the dice task
and executive tests while other cognitive functions were intact.
A positron emission tomography (PET) examination revealed
hypometabolic zones within the dorsolateral prefrontal cortex
(bilateral) as well as the fusiform and the cingulate gyri. We
proposed that decisions under risk can be made by two differ-
ent but interacting ways (Brand et al., 2006). A cognitive way
to select an option relies on explicit knowledge about proba-
bilities and consequences as well as knowledge about the risk
of an option, which is the combination of the probability and
amount of gain/loss. In addition to this cognitive way, we sug-
gest that decisions under risk – as other kinds of decisions,
such as decisions under ambiguity – can also be made on the
basis of feedback from previous decisions. (The role of feed-
back use in decisions under ambiguity according to the somatic
marker hypothesis is described in Bechara, 2001, 2005; Bechara,
Damasio, Tranel, & Damasio, 1997; Bechara, Damasio, Tranel,
& Damasio, 2005; but see also the statements on SCRs measures
and decision-making above.) In decisions under risk, feedback
from previous trials may as well guide the decision-making pro-
cess: If someone chooses a risky option and receives punishment,
additional to his or her prior knowledge about probable nega-
tive consequences, the feedback can be used to reconsider the
current strategy or to more explicitly observe the rules. Under
risk conditions, advantageous performance, i.e. decisions with
maximal rewards, is believed to rely on both logical strate-
gies based on explicit knowledge and the use of feedbacks
from previous decisions. Therefore, structures of both fronto-
striatal loops, the cognitive and the limbic loop (see description
above), are thought to be involved in decision-making under risk
conditions.
So far, the potential role of the amygdala for decisions under
risk is not clear. To our knowledge, this is the first study that aims
to reveal differential contributions of the amygdala to decisions
under ambiguity and decisions under risk in patients with selec-
1307
tive amygdala damage and recording SCRs during performing
both tasks. As mentioned above, gambling task performance
is reported to co-vary with SCRs as a marker of emotional
reactivity. As the amygdala is crucial for processing emotional
stimuli and initiating arousal changes (Phelps, 2006; Phelps &
LeDoux, 2005), we hypothesise – in accordance with findings of
Bechara et al. (1999) – that in our amygdala-damaged patients
generating SCRs is reduced during gambling task performance.
While SCRs have not previously been measured during dice task
performance, behavioural studies revealed that processing feed-
backs from previous trials contribute importantly to successful
dice task performance. As feedback processing seems to rely on
amygdaloid activation and corresponding generation of SCRs
(Bechara et al., 1999), amygdala-damaged patients are hypoth-
esised to demonstrate lower reactions on emotional feedback
and reduced SCRs during dice task performance.
The healthy subjects should generate higher SCRs preced-
ing disadvantageous relative to advantageous decisions in the
gambling task, as revealed in most of the previous studies
which recorded SCRs. In the dice task, healthy subjects are
hypothesised to primarily show higher SCRs for risky relative
to the non-risky decisions in the feedback phase, when gains
or losses are displayed. Anticipatory SCRs should not differ
between risky and non-risky choices, because we believe that
the upcoming decision is based on reflecting about the prob-
abilities and the amount of rewards/punishments more than on
anticipatory reactions that “alert” the individual for the potential
punishment.
On the behavioural level, we hypothesise that in the patients
with UWD, performance in the dice task is less severely affected
than in the gambling task because in the dice task it is possi-
ble to decide on the basis of cognitive strategies from the very
beginning of the task, as mentioned above. More precise, we
hypothesise that in UWD patients one of the suggested ways
to solve the dice task is affected (feedback use and generating
SCRs as markers for emotional processing of the feedbacks)
while the other way (evaluating the options on the basis of their
probabilities and amounts of gains and losses) is intact, resulting
in an overall moderately reduced performance compared with
healthy subjects, who should use both suggested ways to solve
the task.
2. Subjects and methods
2.1. Subjects
We examined three patients suffering from Urbach-Wiethe disease (UWD),
a rare autosomal recessive genetic syndrome producing bilateral calcifications
within the anterior section of the medial temporal lobe, primarily affecting the
amygdaloid complex (see Newton, Rosenberg, Lampert, & O’Brien, 1971; Staut
& Naidich, 1998). One patient (AF) came from Germany, one from Austria (RB)
and one from The Netherlands (WT). The sociodemographic characteristics of
the patients are shown in Table 1. In all three patients, selective bilateral amyg-
dala damage was diagnosed using magnetic resonance imaging (MRI). The
diagnosis of amygdala damage was confirmed by two independent physicians,
who were blind to the clinical presentation of the patients. Potential laterality in
amygdala damage was not reported by the physicians. In patient RB, selective
amygdala calcification was extensive, however, his MRI also revealed slight cal-
cifications in the lentiform nucleus and the uncinate gyrus. In patient WT, the
1308 M. Brand et al. / Neuropsychologia 45 (2007) 1305–1317
Table 1
Sociodemographic characteristics of the UWD patients and the healthy comparison subjects
Patient AF Patient RB
Age (years) 17 38
Gender Female Male
Education (years) 9 13
uncinate and parahippocampal gyri were also slightly affected. In patient AF, no
other signs of calcifications were found beyond the damage of the amygdaloid
complex. In all three patients the main pathology was a clearly visible calci-
fication centred on the amygdaloid complex bilaterally and additional signs,
if present, were reported as non-substantial. All patients had a long-standing
diagnosis of UWD as evidenced by the standard cardinal symptoms (most promi-
nently verrucous nodules and a very hoarse voice in all three patients). In patient
AF, UWD was diagnosed at the age of 6 years, in RB at the age of 4 years and
in WT at the age of 3 years, but symptoms of hoarseness, similarly severe in all
three patients, were existent from the day of birth. In all three patients, the UWD
diagnosis was confirmed by a number of doctors (four as a minimum in AF and
up to eight in WT) prior to our investigation. In patient WT, the symptom of
verrucous nodules was more pronounced than in the other two cases; affecting
his eyes, hands, and elbows, whereas in patients AF and RB, only hands and
elbows were affected.
In addition to the patients we examined 20 healthy comparison subjects with
the experimental tasks (gambling task and dice task, see description below). The
comparison group consisted of individuals matched for age, gender, and educa-
tion and of additional subjects representing normal population. The sociodemo-
graphic characteristics of the comparison group are also summarised in Table 1.
2.2. Neuropsychological test battery and psychological
assessment
The neuropsychological test battery comprised standardised tests for ver-
bal and non-verbal intelligence, anterograde memory, attention, information
processing, and executive functions (see Table 2). Psychiatric symptoms were
assessed by standardised questionnaires: Beck depression inventory (BDI;
Hautzinger, Bailer, Worall, & Keller, 1995) and the Symptom Check List (SCL-
90-R; Franke, 2002).
2.3. Decision-making tasks
For decision-making we administered two tasks: the computerised version of
the Iowa Gambling Task (“gambling task”) (Bechara, Tranel, et al., 2000) and the
computerised Game of Dice Task (“dice task”) (see Brand, Fujiwara et al., 2005).
The gambling task involves four decks of cards, decks A, B, C and D from which
participants have to choose one card in each of 100 trials. Each time a subject
selects a card, a specific amount of fictitious money is awarded. However, at
certain times, losses of different fixed amounts occur. Two of the decks of cards,
decks C and D, are considered advantageous, as they result in small immediate
gains, but also very small losses and will therefore bring in more money than they
take in the long run. The other two decks, decks A and B, produce high immediate
gains but will take more money than they give due to very high losses at certain
times. Therefore, these decks are considered to be disadvantageous (see detailed
instructions for the gambling task in Bechara, Tranel, et al., 2000). In summary,
participants are told the goal of the task is to gain as much money as possible
and to avoid losing money. They are also informed they can switch between
decks at any time. It is not disclosed to the participants that the task consists of
100 card selections and which card decks are advantageous or disadvantageous,
but they are informed that some of the decks are better than others. Thus, the
exact rules for gains and losses and winning probabilities are not explicit to
the subjects. Rather, they have to learn to avoid the disadvantageous decks and
to prefer the advantageous decks by remembering and using their feedback
from previous trials. Accordingly, the gambling task measures decisions under
ambiguity, at least at a stage before participants have implicitly or explicitly both
learned and implemented maximally advantageous strategies. For the analysis of
gambling task performance we calculated a netscore by subtracting the number
Patient WT Comparison group
50 M = 30.35 (S.D. = 13.06)
Male 11 female and 9 male
10 M = 12.50 (S.D. = 1.23)
of disadvantageous choices (decks A and B) from the number of advantageous
choices (decks C and D). Thus, higher netscores indicate better performance on
the task and negative netscores represent lower task performance, i.e. selection
of more disadvantageous than advantageous decks.
In contrast to the gambling task, the dice task offers explicit rules for gains
and losses as well as obvious winning probabilities associated with each option.
In the computerised dice task, subjects are instructed to maximise a fictitious
starting capital by choosing one of four different alternatives in each of 18 trials
in which one single virtual dice is thrown. The options differ in their probability
to yield a reward (1:6 to 4:6). For options with low winning probabilities (less
than 50%), the gains and losses are high (D 1000 for one single number and D 500
for a combination of two numbers). For options with high winning probabilities
(50% and higher), the gains and losses are moderate to low (D 200 for a com-
bination of three numbers and D 100 for a combination of four numbers). The
contingencies are displayed in Fig. 1. The rules and amounts of gains and losses
are explicitly described in the test instruction and are permanently visualised on
the screen. Subjects are also informed about the total of 18 decisions within the
game. After each throw, the gain or loss (depending on congruence or incon-
gruence between the selected number(s) and the thrown number) is indicated
on the screen. The computer also indicates the participant’s current financial
balance, as well as the number of remaining dice throws. As outlined above,
the dice task represents decision situations with explicit gains and losses as well
as probabilities. Therefore, this task measures decisions under risk, when the
outcome of an option is explicitly defined by probabilities (see Bechara, 2004).
A more detailed task description can be found elsewhere (Brand, Fujiwara et al.,
2005; Brand, Kalbe et al., 2005). When analysing dice task performance, two of
the four possible alternatives (combination of three numbers and combination of
four numbers) are classified ‘advantageous’ since they have a winning probabil-
ity of 50% and higher and are associated with low gains but also low losses (see
Fig. 1). Therefore, they most likely result in a positive balance long-term. The
remaining two alternatives (a single number and a combination of two numbers)
are classified as ‘disadvantageous’ as they have a winning probability of lower
than 50% and result in high gains but also high losses. In order to compare
performance on the dice task and on the gambling task, we also calculated a
total netscore for the dice task by subtracting the number of disadvantageous
choices from that of advantageous selections [(sum of choices of three and four
numbers) − (sum of choices of one number and two numbers)].
The order of gambling task and dice task administration was randomised.
In the comparison group, 10 subjects performed the gambling task first and 10
subjects performed the dice task first. As described in previous studies (e.g.,
Brand, Recknor, Grabenhorst, & Bechara, in press), there was no effect of task
order in our healthy individuals (gambling task: t = 0.078, p = .939; dice task:
t = 0.883, p = .389). On a descriptive level, there was also no effect of task order
in our three patients. Two patients performed the gambling task before the dice
task (RB and WT) and one patient had the reverse order (AF). As presented in
the results section, one of the two patients, who started with the gambling task
had lower scores in this task than the comparison group (RB), the other patient
(WT) was within normal range. On the dice task, also one of these patients (WT)
performed lower than the healthy subjects and the other patient (RB) performed
normally. Patient AF, who performed the dice task first showed lower scores in
both tasks compared with the healthy subjects.
2.4. Skin conductance responses (SCRs)
During performance of the gambling task and the dice task, we recorded
electrodermal activity as a measure of emotional reactivity. The skin conduc-
tance responses (SCRs) were registered via two Ag/AgCl electrodes attached
to the thenar and hypothenar areas of the subjects’ non-dominant hand. During
 M. Brand et al. / Neuropsychologia 45 (2007) 1305–1317 1309

Table 2
Neuropsychological profile of the UWD patients (references for the mentioned tests can be found elsewhere, e.g., in Lezak, Howieson, & Loring, 2004; Spreen &
Strauss, 1998)
Domain/tests Max AF RB WT

RS P RS P RS P

Information processing and executive functions

Nelson’s Modified Card Sorting Test (MCST)
Categories 6 6 69 5 34 1 0
Non-perseverative errors 4 58 6 48 2 73
Perseverations 1 78 5 50 33 0
Trail Making Test A s 22 70 26 50 59 10
Trail Making Test B s 64 53 55 55 90 10
Word Colour Interference Test
Reading the word s 41 10 27 69 37 50
Naming the colour s 75 8 45 50 63 16
Interference trial s 100 27 86 50 104 42
Interference—naming s 25 41 41
Controlled Oral Word
Association (FAS) Test 28 10 46 50 20 <10
Verbal Fluency “Animals” 15 5 24 48 16 9
Tower of Hanoi (three discs)
Moves 11 8 9
Time s 63 21 57
Errors 0 0 3
Attention
Selective Attention Test
(d2) Total errors 375 15.9 522 99.2 293 30.9
Intelligence
Subtest verbal reasoning (LPS) 40 23 42 34 98 18 31
Estimated IQ 98 130 94
Visuo-Constructive Abilities
Rey-Osterrieth-Figure
Copy 36 32 16 33 50 34 76.5
Anterograde Memory
Rey-Osterrieth-Figure
Delayed recall 36 18 27 23.5 76.5 15.5 50
Affective Words Test
Delayed free recall 15 4 <15 2 <5 1 <1
Delayed recognition 15 14 66 15 82 14 66
Malingering
Test of Memory Malingering
Trial 1 50 49 73 50 84 50 84
Trial 2 50 49 16 50 66 50 66
Retention 50 50 66 50 66 49 16
Theory of Mind and Emotional Processing
ToM Eyes Test 24 15 12 19 62 12 1
Florida Affective Battery
Facial affect discrimination % 87.71 27 95 73 80 18
Facial affect naming % 93.3 42 90 50 65 4
Facial affect attribution % 100 73 100 69 80 10
Affective prosody discrimination % 100 62 100 66 93.3 2
Affective prosody naming % 93.3 24 75 2 64.3 0
Depression
Beck’s Depression Inventory 63 4 0 0
Psychological-psychiatric symptoms
SCL-90-R
Somatisation t-Scores 52 42
Obsessive-compulsive 55 40
Interpersonal sensitivity 35 38
Depression 45 36
Anxiety 43 40
1310 M. Brand et al. / Neuropsychologia 45 (2007) 1305–1317

Table 2 (Continued )
Domain/tests Max AF RB WT

RS P RS P RS P

Anger-hostility 46 41
Phobic anxiety 43 45
Paranoid ideation 46 41
Psychoticism 39 43
Global severity index 45 41

RS = raw scores; P = percentiles; Max = maximum; s = seconds; % = scores given in percent; LPS = German intelligence test (Leistungsprüfsystem); t-scores between
40 and 60 represent scores within the range of one standard deviation of the norm group’s scores; italicised scores indicate impaired performance. Note: The Affective
Words Test is an experimental task based on the material described in Brand et al. (2003). Affective words have to be evaluated regarding their valence (negative,
neutral or positive). After a delay of 15 min, these words have to be freely recalled and recognised. Participants are not informed that they have to memorise the
words. Therefore, the judgment trial acts as an implicit learning trial for the delayed recall of the words. Percentiles for delayed recall and the recognition of the
words are based on a sample of 30 healthy individuals.

administration of both tasks subjects were seated in a comfortable chair in
front of a computer screen while SCRs were continuously recorded with a
PhysioModul (med-NATIC, Munich) system. The acquired data were stored
and analysed on a computer with a LDI700-system and -software.
In the gambling task, the time window for the reward and punishment SCRs
was 5 s after choosing a card by clicking on its picture on the computer screen
(measuring started immediately after the choice). In the dice task, due to a 2.5 s
delay between selection and feedback delivery, the time window for the reward
and punishment SCRs was 5 s after feedback had been delivered (measuring
started immediately after feedback was given, i.e., 2.5 s after the choice). In both
tasks SCRs generated after the end of the time window for reward/punishment
SCRs and before the selection on the next trial were analysed as anticipatory
SCRs. For analysis of reward/punishment SCRs as well as anticipatory SCRs
the amplitude of each SCR was measured. The inter-trial intervals in both tasks
were set at 6 s to allow for psychophysiological recordings (Bechara et al., 1999).
This interval was chosen in order to avoid potential overlaps between feedback
reactions and anticipatory reactions on the upcoming trial and to allow the skin
conductance level to return to a baseline. Trials with artifacts due to movement
or breathing were excluded from the analysis. Individual SCRs were standard-
ised according to Lykken and Venables (1971). All SCR measures are given in
␮Siemens (␮S).
2.5. Statistical analyses
For the main behavioural data, we present raw scores of each patient sepa-
rately and indicate whether or not the performance is within 95% of the healthy
comparison subjects’ performance (=1.64 S.D.s). In the analysis of the compar-
ison group’s SCR measures, we additionally conducted t-tests for dependent
samples in order to compare SCRs for disadvantageous and advantageous deci-
sions in the gambling task or risky and non-risky decisions in the dice task,
respectively. Results were corrected for multiple comparisons (Bonferroni) when
appropriate. We also calculated Pearson correlations in order to analyse the
potential relationships between generating SCRs and gambling and dice task
performance in the healthy subjects. The statistical analyses were carried out
with SPSS version 12.0 for Windows (Chicago, SPSS Inc.).
3. Results
3.1. Results in the neuropsychological test battery
The results in the neuropsychological test battery are shown
in Table 2. As indicated by the percentiles, two patients (AF

Fig. 1. Summary of the options in the dice task. Participants can choose one single number (one to six; six options altogether) associated with a gain of D 1000 if the
selected number is thrown and with a loss of D 1000 if another number than the selected is thrown. Participants can also choose a combination of two numbers (three
different options). If one of the numbers included in the selected option is thrown, the participants wins D 500. In case that another number is thrown, the participant
looses D 500. The combinations of three numbers together are linked to D 200 gain/loss and the most conservative combinations of four numbers are related to D 100
gain/loss. The options of one number and the options consisting of two numbers are analysed as “risky” because the have a winning probability of less than 50%
but are associated with high gains/losses. The options of combinations of three numbers and four numbers are analysed as “non-risky” because they have a winning
probability of 50% and higher, most likely leading to a positive balance in the long run.
 M. Brand et al. / Neuropsychologia 45 (2007) 1305–1317
and WT) were impaired in at least two out of the five tasks
assessing executive functions and other cognitive domains while
the remaining patient was unimpaired across all five executive
tasks. The psychiatric screenings did not show clinically rele-
vant symptoms of depression or other psychiatric disorders in
any patient. Unfortunately, we did not have a Dutch version
of the Symptom Check List. Therefore, with patient WT this
check list was not given as a paper-pencil questionnaire, but as
an interview without the detailed scoring system (only yes/no
answers). Accordingly, we did not calculate the t-scores. In our
ad-hoc interview, which was based on the items of the Symp-
tom Check List, patient WT did not report any of the symptoms.
On this basis, we also exclude psychiatric symptoms in patient
WT.
3.2. Results in the decision-making tasks
The comparison group’s mean gambling task total netscore
was M = 27.00 (S.D. = 22.26). On a single case level, perfor-
mance on the gambling task of two patients (AF and RB) was
outside of 1.64 S.D.s (=95% interval of the variance) of the
comparison group’s mean performance (total netscore of patient
AF = −10; patient RB = −38; interval of 1.64 S.D.s in the com-
parison group = −9.5 to 63.5). The third patient (WT) showed a
profile in the gambling task indicating random decisions with a
netscore of 2. His result shows that he selected advantageous and
disadvantageous card decks with an equal frequency without any
preference. Fig. 2 illustrates performance of the three patients
and the mean gambling task performance of the comparison
group. In order to make the behavioural results comparable to
those presented in the section on psychophysiological recording,
Fig. 2 includes the total number of advantageous and disadvan-
Fig. 2. Performance in the gambling task divided into the frequency of choices
from the advantageous and from the disadvantageous card decks. Results of
the three UWD patients are shown separately and contrasted with the mean
performance of the comparison group (CG). Error bars in the comparison group
indicate 1.64 S.D.s (=95% variance of the comparison group’s performance).
1311
tageous decisions. Note that the gambling task netscore is the
subtraction of the number of disadvantageous decisions from
the number of advantageous choices.
In the dice task, the comparison subjects had a mean netscore
of M = 10.75 (S.D. = 7.30). On a single case level, two patients
(AF and WT) performed outside of 95% of the comparison
subjects’ variance (dice task netscore of patient AF = −10;
patient WT = −6; interval of 1.64 S.D.s in the comparison
group = −1.22 to 22.72). These two patients also used the neg-
ative feedback from previous risky decisions less frequently
than the comparison subjects. Using the negative feedback was
defined as the following: If subjects had chosen a risky alterna-
tive and received a negative feedback (loss of money) in the pre-
vious trial and then chose a non-risky alternative in the following
trial, this is rated as ‘used negative feedback for a decision-shift
to a non-risky alternative’. Patient AF used 22.22% of the neg-
ative feedback for a decision shift and WT used 27.27%. The
comparison group on average used 85.60% (S.D. = 21.11) of the
negative feedback following a risky decision. The scores of feed-
back use of patients AF and WT were outside of the 95% inter-
val of the comparison group’s variance (bottom score = 50.98).
In contrast, patient RB performed normally on the dice task
(netscore = 12) and used 100% of the negative feedback follow-
ing a risky decision. Interestingly, patients AF and WT were
also the patients with compromised executive functions in at
least two out of the five measures administered, whereas RB
was unimpaired in formal neuropsychological testing (compare
Table 2). In Fig. 3, the number of advantageous and disadvanta-
geous decisions in the dice task are presented separately for each
patient and compared with the mean performance and 1.64 S.D.s
of the comparison subjects.
Fig. 3. Performance in the dice task divided into the frequency of risky and
non-risky choices. Results of the three UWD patients are shown separately and
contrasted with mean performance of the comparison group (CG). Error bars
in the comparison group indicate 1.64 S.D.s (=95% variance of the comparison
group’s performance).
1312 M. Brand et al. / Neuropsychologia 45 (2007) 1305–1317
Fig. 4. Skin conductance responses (SCRs) of the three patients and the com-
parison group during performance of the gambling task. SCRs are divided
into anticipatory and feedback SCRs before and after choosing a card from
an advantageous and disadvantageous card deck, respectively. Error bars in the
comparison group indicate 1.64 S.D.s (=95% variance of the comparison group’s
SCRs).
3.3. Results in SCRs measurements during gambling task
and dice task performance
We analysed SCRs in two phases of both the gambling
task and the dice task: before a decision was made (anticipa-
tory SCRs) and after receiving the feedback (feedback SCRs).
Both SCR measures were calculated separately for advanta-
geous and disadvantageous decisions in the gambling task
and the dice task. The comparison group’s mean anticipatory
SCRs for disadvantageous decisions in the gambling task were
M = 0.216 ␮S (S.D. = 0.109) and M = 0.143 ␮S (S.D. = 0.098)
for advantageous decisions, respectively. The mean feedback
SCRs of the comparison group after choosing a disadvantageous
card deck were M = 0.269 ␮S (S.D. = 0.125) and M = 0.189 ␮S
(S.D. = 0.119) after selecting an advantageous alternative. Fig. 4
shows the SCRs for advantageous and disadvantageous deci-
sions in the gambling task for the comparison group and the
three patients. It can be seen that for disadvantageous decisions,
both the anticipatory and the feedback SCRs are lower than 95%
of the comparison group’s variance in all three patients. In con-
trast, one patient (RB) produced SCRs within a 95% interval
of the comparison group’s variance before and after choosing
an advantageous alternative, while the other two patients also
produced reduced SCRs (outside of the 95% interval of the com-
parison group’s variance).
In the dice task, the healthy individuals generated the follow-
ing SCRs: anticipatory SCR for risky decisions: M = 4.27 ␮S
(S.D. = 0.159), anticipatory SCR for non-risky decisions:
M = 0.321 ␮S (S.D. = 0.212), feedback SCR for risky deci-
Fig. 5. Skin conductance responses (SCRs) of the three patients and the compari-
son group during performance of the dice task. SCRs are divided into anticipatory
and feedback SCRs before and after choosing risky or non-risky options in the
dice task, respectively. Error bars in the comparison group indicate 1.64 S.D.s
(=95% variance of the comparison group’s SCRs).
sions: M = 0.416 ␮S (S.D. = 0.164), and feedback SCR for non-
risky decisions: M = 0.272 ␮S (S.D. = 0.176). The SCRs of each
patient compared with the mean of the comparison group and
95% interval of variance in the comparison group are displayed
in Fig. 5. All three patients generated reduced SCRs before
selecting a risky alternative (outside of the 95% interval of the
comparison group’s variance) (see Fig. 5). The feedback SCRs
for risky decisions were reduced in two patients (AF and WT).
In the comparison group the SCRs for disadvantageous
and advantageous decisions in the gambling task differed
significantly (p-value after correction for multiple compar-
isons = .0125) in both anticipatory (t = 6.76, p < .001) and feed-
back phases (t = 4.24, p < .001). In the dice task, the SCRs for
risky and non-risky decisions were significantly different for the
feedback phases (t = 4.11, p = .001) but not for the anticipatory
phases (t = 2.17, p = .046). In addition, the netscore of the dice
task was correlated with anticipatory SCRs for risky decisions
(r = .550, p = .027). Other correlations between SCRs and dice
task or gambling task performance failed to reach significance
(they were between r = .20 and r = .45).
4. Discussion
The results of our study confirm the main hypothesis that
patients with amygdala damage show reduced performance in
both, decisions under ambiguity (measured with the gambling
task) and decisions under risk (measured with the dice task).
The observed lower gambling task performance in the patients
replicate a finding by Bechara et al. (1999), reporting deficient
decision-making in the gambling task in their sample of five
 M. Brand et al. / Neuropsychologia 45 (2007) 1305–1317
patients with amygdala damage. Our patients with amygdala
damage generated reduced SCRs during the anticipatory phases
as well as after receiving feedback from previous trials in the
gambling task. This pattern is also in accordance with the pre-
viously mentioned study of Bechara et al. (1999) and can be
interpreted in the light of the somatic marker theory (Damasio,
1994, 1996), as Bechara et al. (1999) argued that compromised
decision-making in amygdala-damaged patients is “an indirect
consequence of the role of the amygdala in attaching affective
attributes to stimuli”.
As mentioned in Section 1, there is some evidence for a
relationship between good gambling task performance and gen-
erating SCRs, primarily before picking a card from the dis-
advantageous decks (Bechara et al., 1997; Crone et al., 2004;
Denburg, Recknor, Bechara, & Tranel, 2006; Hinson et al.,
2002). In addition, feedback SCRs are also discussed as influ-
encing task performance, as they potentially reflect appraisal
of the decision. Findings in animal studies and neuroimaging
investigations in both healthy volunteers and brain damaged
patients have demonstrated that the amygdala is a structure cru-
cially involved in emotional implicit learning beyond simple fear
conditioning processes (see the recent articles of Phelps, 2006;
Phelps & LeDoux, 2005). Accordingly, patients with amyg-
dala damage may be deficient in implicitly learning to avoid
the disadvantageous decks and to establish strategies in order
to maximise rewards using previous feedback. A reason for this
deficit may be that the punishments (lost money) do not evoke
significant somatic states, as reflected by reduced SCRs that can
be used in future decision situations as discussed by Bechara et
al. (1999) (see also Bechara, 2004; Bechara et al., 2003, 2005).
However, the relationship between generating SCRs and amyg-
dala functioning is still a topic of debate. While studies with
functional imaging techniques revealed such a correlation (e.g.,
Furmark, Fischer, Wik, Larsson, & Fredrikson, 1997; Williams
et al., 2005), and studies with amaygdala-damaged patients
reported altered SCRs in these patients (see above), there are
also case-reports which indicate that generating SCRs (e.g., in
a conditioning paradigm) can be normal in patients with amyg-
dala damage (Tranel & Damasio, 1989; Tranel & Hyman, 1990).
In summary, most of the studies, however, indicate that there
is at least an unspecific general association between amygdala
functioning and generating physiological responses to emotional
stimuli (changes in heart rate, blood pressure, electrodermal
activity), due to the connections between amygdala and brain-
stem nuclei involved in regulating autonomic responses (see
Davis & Whalen, 2001).
The main aim of our investigation was, however, to anal-
yse the possible role of the amygdala in decisions under risk.
When analysing dice task performance on a single-case level,
two out of the three patients with amygdala damage had a neg-
ative netscore in the dice task and performed outside of 95%
of the comparison group’s variance. These two patients also
had executive deficits in at least two of the administered execu-
tive tasks, while the remaining patient showed normal executive
functioning and dice task performance compared to healthy indi-
viduals. This result is in accordance with previous studies that
focused on the relationships between dice task performance and
1313
executive functions (e.g., Brand, Fujiwara et al., 2005; Brand,
Kalbe et al., 2005; Brand, Labudda et al., 2004). As described
in the former articles, we suggested that decisions under risk
can be made using two different but interacting ways: an emo-
tional and a cognitive way. However, we proposed that good
performance in tasks assessing decisions under risk requires the
integration of both the use of feedback following previous trials
(as described for decisions under ambiguity; see Bechara, 2001,
2005; Bechara, Damasio, et al., 2000; Bechara et al., 1997, 1999,
2003, 2005; Bechara & Van Der Linden, 2005; Tranel, Bechara,
& Denburg, 2002) and, in addition, cognitive functions such as
categorisation of the options offered by the decision situation,
recognition of probabilities and the amount of gains and losses
and so on. Evidence for the importance of feedback comes from
the two patients with lower scores in the dice task, who also
used the negative feedback following a risky decision less fre-
quently in order to shift to a non-risky option in the upcoming
decision. In addition, in the healthy subjects the feedback SCRs
differed between risky and non-risky decisions, but the antic-
ipatory SCRs preceding risky and non-risky decisions did not
differ significantly. Interestingly, dice task performance was not
correlated with the level of feedback SCRs after risky decisions
but with the anticipatory SCRs before choosing a risky option.
This was not expected as we thought that before making a deci-
sion in the dice task, cognitive processes would be predominant
and that “warning signals” from the periphery would not play
a major role. Further studies should investigate SCR correlates
of dice task performance using a larger sample of individuals in
order to assess the potential role of anticipatory and feedback
SCRs for dice task performance in more detail.
However, the current results give support for the assumption
that decision-making under risk conditions involves both the
integration of feedback from previous trials as well as strategic
aspects. In case of deteriorations of both components, deficits in
such decisions are more likely than in the case of impairments
in only one of these components (e.g., only deficits in emotional
reactivity or strategy application, respectively). This assump-
tion is consistent with results of previous studies, as mentioned
above. More specifically, the descriptive subgroup analysis of
dice task performance in patients with Morbus Parkinson (see
Brand, Labudda et al., 2004) indicated that patients who were
neither deficient relative to healthy control subjects in feedback
processing (as measured by the number of negative feedbacks
following a risky decision in the dice task that triggered a shift
to the non-risky alternatives) nor impaired in executive func-
tioning performed well on the dice task. In contrast, patients
with executive dysfunctions but intact feedback use as well as
those with selective deficits in the use of feedback but intact
executive functions performed worse in the dice task. How-
ever, the strongest impairments in the dice task were shown in
patients with executive deficits and reduced feedback use. In the
present study, a similar association–dissociation can be found.
The two patients who had at least reduced executive function-
ing in some of the administered executive tasks, were strongly
reduced in dice task performance, while the other patient per-
formed normally. This result further supports the assumption of
the importance of executive functions – primarily categoriza-
1314 M. Brand et al. / Neuropsychologia 45 (2007) 1305–1317
tion, set-shifting and strategy application – for decision-making
in a situation with explicit rules for rewards and punishments
and obvious winning probabilities. In addition, the two patients
with low dice task performance also showed reduced feedback
processing in this task. This result is also in accordance with
the aforementioned studies on dice task performance with dif-
ferent patient groups. Nevertheless, one has to keep in mind that
these two patients (AF and WT) were also different from the
patient with normal dice task performance (RB) in a few other
aspects. They also had problems in theory-of-mind functions and
emotional processing, which could contribute to their difficul-
ties in decision-making, e.g., in processing negative feedback.
In addition, they had lower intelligence scores (measured by
logical reasoning) which could be interpreted that their reduced
decision-making performance reflects general intellectual con-
straints. However, in most of the other tasks they performed
normally (e.g., on attention, anterograde memory, problem solv-
ing) and their intellectual abilities were within the normal range.
Thus, it seems unlikely that they were unable to understand
or remember the task instructions. Furthermore, previous stud-
ies on dice task performance in healthy subjects and different
patient groups (e.g., Brand, Fujiwara et al., 2005; Brand, Kalbe et
al., 2005; Brand, Labudda et al., 2004), revealed no correlation
between dice task performance and measures of intelligence.
Therefore, we believe that the poor performance on the dice
task of patients AF and WT was not generally influenced by
their intelligence level that was normal but lower than that of
patient RB. One has also to notice that patient RB, who had an
average IQ of 130, was one of the two patients who performed
outside of 95% of the comparison group’s variance in the gam-
bling task. This result emphasises that decision-making is not
primarily influenced by intelligence and is in line with the study
of Brand et al. (in press) which revealed that in healthy subjects,
only executive functions were predictors of gambling task and
dice task performance but not intelligence, age, gender or other
sociodemographic variables.
One may consider that the additional signs of calcifications
in the uncinate gyrus in two of the three patients, as described
in the methods section, were accountable for the differences
in decision-making and executive functions across the three
patients. However, patient AF scored lower than the comparison
group on both the gambling task and the dice task and had impair-
ments in two of the executive measures, yet she was the patient
without any further signs of calcifications beyond the amygdala
degeneration. In contrast, patient RB, who had slight signs of
calcifications in the lentiform nucleus as well as in the uncinate
gyrus performed well on the dice task and had no deficits across
the executive tests administered, high IQ and overall relatively
good performance on most of the neuropsychological functions
assessed.
The question remains, why patients with selective amygdala
damage, as our patients with UWD, can have executive dysfunc-
tions, even though only some aspects of executive functions were
affected in two patients (AF and WT) and their impairments were
heterogeneous across the executive tasks administered. Both
patients were deficient in verbal fluency, but only patient WT was
additionally impaired in the Modified Card Sorting Test. Patient
AF showed increased interference susceptibility in the Word
Colour Interference Test. In the same task, patient WT was not
clinically impaired but his percentile was 16, indicating perfor-
mance within the lower interval of normal scores (percentiles of
15 and below are interpreted as impaired). This pattern of hetero-
geneous executive task performance in our patients might reflect
the variety of subcomponents of executive functions which can
be individually affected. Due to the fact that we do not have any
information about potential functional changes of frontal brain
areas in our sample of patients, a comprehensive explanation of
the patients’ profile is problematic, as the specific subcompo-
nents of executive functioning most likely have both common
and divergent neural correlates. The result that UWD patients
can have difficulties in executive tasks is nevertheless interesting
by itself, since the question of whether or not executive dysfunc-
tions can occur in the course of UWD is still a topic of debate. In
most of the previous studies on neuropsychological impairments
following amygdala damage, executive functions were consid-
ered to be more or less intact. For instance, in the study of Shaw
et al. (2004) as well as in the single-case study of Fine, Lumsden,
and Blair (2001), deficits in social cognition or “theory of mind”
in patients with amygdala lesions were dissociated from exec-
utive functioning. Additionally, the two patients with UWD,
studied by Markowitsch et al. (1994), did not show abnormalities
in executive tasks performance. Further evidence for generally
intact executive functioning in patients with UWD comes from
the study of Siebert, Markowitsch, and Bartel (2003), in which
the so far largest sample of UWD patients (n = 10) has been
neuropsychologically examined. The authors did not find gen-
eral executive impairments, but compared with the comparison
subjects some of the patients showed reduced performance in
the Trail Making Test B. However, as the focus of the study by
Siebert and colleagues was the differential analysis of emotional
processing and emotional memory, executive functioning was
not examined extensively (only with two tasks). Therefore, it is
difficult to conclude that UWD patients are completely unim-
paired in executive functions on the basis of previous results.
In contrast, the single-case study of a UWD patient (SM) by
Tranel and Hyman (1990) indicates that executive functions
can be disturbed in the cause of the disease beyond memory
impairments and deficits in social behaviour. The patient SM
was severely deficient in several tasks assessing category forma-
tion, cognitive flexibility, set-shifting, and verbal fluency. Even
though functional neuroimaging techniques (e.g., PET or fMRI)
have not been conducted and therefore functional brain changes
outside the amygdala cannot be excluded definitely, Tranel and
Hyman concluded that impairments in executive control and
social behaviour are due to the selective mineralisation of the
bilateral amygdaloid complex. This is also in accordance with
the assumption that the amygdala acts as a modulator of (emo-
tional) reactions which are probably also involved in controlling
behaviour and other executive functions (Phelps, 2006).
Given that the amygdala is directly and indirectly inter-
connected with various subcortical and cortical structures (see
LeDoux, 2000; Nieuwenhuys, 1996; Sarter & Markowitsch,
1985), one might assume that functional alterations of these
connections due to amygdaloid mineralisation can result in
 M. Brand et al. / Neuropsychologia 45 (2007) 1305–1317
functional (especially neurochemical) abnormalities in differ-
ent brain regions and associated neuropsychological changes.
Although in monkeys the amygdala is directly connected with
the dorsolateral prefrontal cortex (see Aggleton & Saunders,
2000), such a direct connection has not been described in humans
(see Phelps, 2006). However, there are several indirect con-
nections between the amygdala and the dorsolateral prefrontal
cortex. One of those connections is the ventral amygdalofugal
tract, connecting the amygdala with the mediodorsal thalamus,
which in turn is reciprocally directly and indirectly connected
with the subcallosal area of the basal forebrain, the dorsolateral
prefrontal cortex and the anterior cingulate gyrus. Functional or
structural pathology of the ventral amygdalofugal tract may be
responsible for the development of executive control decrease.
Thus, executive dysfunctions might occur as an indirect conse-
quence of amygdala damage in the course of UWD. However,
studies that investigate functional brain abnormalities using neu-
roimaging techniques (e.g., glucose utilisation using PET) in
patients with UWD are needed in order to reveal possible frontal
abnormalities in UWD. The duration and life-time onset of
amygdala degeneration could also play a role for both decision-
making difficulties and executive dysfunctions in patients with
UWD. If amygdala damage occurred early in life, then frontal
regions do not receive appropriate input from the amygdala. As a
consequence, patients may become impaired in developing ade-
quate high-level cognitive strategies (see Hamann et al., 1996).
Accordingly, patients with early amygdala damage should be
reduced in both gambling and dice task performance as well as
executive functioning. In contrast, patients with late amygdala
damage could have developed normal frontal brain functions and
thus be deficient relative to control subjects in the gambling task,
while dice task performance should only be slightly affected due
to intact frontal functions that allowed the possibility to decide
advantageously on the basis of strategies and other cognitive
processes (such as evaluation of winning probabilities and asso-
ciated gains and losses). Unfortunately, we had no information
about the exact time of amygdala degeneration onset. We only
know that all patients had UWD since early childhood, but this
does not necessarily involve that amygdala calcification began at
this point in time. Future studies should address this topic more
extensively.
The second hypothesised important part of the process of
decision-making under risk conditions, as described above, is
the use of feedback from previous trials for current decisions.
In this context, we suggest that emotional reactions on the feed-
back act as biasing signals and influence the upcoming decisions
comparable to their functions in decisions under ambiguity (e.g.,
appraisal of the decision). The reduced SCRs in our patients in
the dice task support this hypothesis. As in the gambling task, all
three UWD patients generated decreased SCRs in the anticipa-
tory phase of the dice task. However, patient RB, who performed
normally in this task, also had reduced SCRs in the anticipatory
phase, but showed normal SCRs in the feedback phase. By con-
trast, the other two patients also generated reduced SCRs in the
feedback phase. Given the results regarding SCRs in the gam-
bling task and the dice task obtained from the within-subjects
comparisons in the healthy individuals, one might assume that
1315
in participants with intact cognitive – especially executive –
functions, the feedback following a decision is used to evalu-
ate whether or not a decision was a “good” decision, but that the
feedback does not necessarily influence the upcoming decision.
In the comparison group, the SCRs in the dice task were higher
for risky decisions than for non-risky only in the feedback phases
but not in the anticipatory phases. This result may indicate that
in healthy subjects, decisions under risk relied more on explicit
information and strategic components, whereas biasing signals
(measured by anticipatory SCRs) played a minor role. Neverthe-
less, emotional reactions to feedback (measured by the feedback
SCRs) may lead to an evaluation of the strategy applied and may
indirectly modify the decision-making process.
5. Conclusion
In summary, we assume that – in contrast to decisions under
ambiguity – decisions under risk rely on both cognitive and emo-
tional components of the decision situation. Decisions under
ambiguity are more related to emotional feedback processing
and generating of biasing signals to direct future decisions. In
patients with selective damage of the amygdala both decisions
under ambiguity and decisions under risk are affected, the latter
being more reduced, if executive functioning is also compro-
mised.
Acknowledgements
We thank Esther Fujiwara (Rotman Research Institute
Toronto, Canada) for helpful comments on this paper. We also
thank Eva Böcker for her assistance with the graphs. Parts of
the work on this study were supported by the German Research
Foundation (BR 2894/1-1).
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