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www.elsevier.com/locate/brainres

Research Report

Neurodynamics of an election

Armando Freitas da Rocha a,b,⁎, Fábio Theoto Rocha a,b ,

Marcelo Nascimento Burattini b , Eduardo Massad b
a
RANI-Research on Natural and Artificial Intelligence, Rua Tenente Ary Aps, 172 Jundiaí, CEP 13207-110, Brazil
b
School of Medicine, University of São Paulo and LIM01-HCFMUSP, Rua Teodoro Sampaio 115, São Paulo, CEP 5405-000, SP, Brazil

A R T I C LE I N FO AB S T R A C T

Article history: Variables influencing decision-making in real settings, as in the case of voting decisions, are
Accepted 21 June 2010 uncontrollable and in many times even unknown to the experimenter. In this case, the
Available online 26 June 2010 experimenter has to study the intention to decide (vote) as close as possible in time to the
moment of the real decision (election day). Here, we investigated the brain activity
Keywords: associated with the voting intention declared 1 week before the election day of the Brazilian
EEG Firearms Control Referendum about prohibiting the commerce of firearms. Two alliances
Decision-making arose in the Congress to run the campaigns for YES (for the prohibition of firearm
Voting commerce) and NO (against the prohibition of firearm commerce) voting. Time constraints
Brain networks imposed by the necessity of studying a reasonable number (here, 32) of voters during a very
ERP short time (5 days) made the EEG the tool of choice for recording the brain activity associated
Distributed processing with voting decision. Recent fMRI and EEG studies have shown decision-making as a process
Benefits due to the enrollment of defined neuronal networks. In this work, a special EEG technique is
Risks applied to study the topology of the voting decision-making networks and is compared to
the results of standard ERP procedures. The results show that voting decision-making
enrolled networks in charge of calculating the benefits and risks of the decision of
prohibiting or allowing firearm commerce and that the topology of such networks was vote-
(i.e., YES/NO-) sensitive.
© 2010 Elsevier B.V. All rights reserved.

1. Introduction most real settings of decision-making, in which the influenc-

New brain imaging techniques have opened the way for the
study of the processes of human decision-making. This kind
of research is usually done under simulated conditions
that allows for substantial control, by the researcher, of the
most important variables influencing the decision-making
(Knutson et al., 2007; Greene et al., 2004; McClure et al., 2004;
Moll and Oliveira-Souza, 2007; Huettel et al., 2006). These
controlled simulated conditions are in sharp contrast with
ing variables are uncontrollable and in many times even
unknown. As a matter of fact, we may consider that some
decision-making processes are always hypothetical (e.g.,
moral judgment) in the sense that there is no way to carry
out experiments in real cases; some other kinds of decisions
(e.g., purchasing) allow simulated (and controlled) experimen-
tation, and, finally, some others (e.g., voting) are always done
in real and uncontrolled environments. In this latter case, the
best that can be done is to study the intention to decide as

⁎ Corresponding author. Rua Tenente Ary Aps, 172, Jundiaí, Brazil, CEP 13207-110. Fax: +55 11 3379 2010.
E-mail addresses: armando@enscer.com.br (A.F. da Rocha), fabio@enscer.com.br (F.T. Rocha), mnburatti@usp.br (M.N. Burattini),
edmassad@usp.br (E. Massad).
URL: http://www.eina.com.br (A.F. da Rocha).

0006-8993/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.brainres.2010.06.046
 Author's personal copy
BR A I N R ES E A RC H 1 3 5 1 ( 2 01 0 ) 1 9 8 – 2 11
close as possible in time to the moment of the real decision.
This is the approach used in this paper that investigates the
brain activity associated with the voting intention declared
1 week before the election day of the Brazilian Firearms
Control Referendum.
The Firearm Commerce Referendum called for a collective
decision about prohibiting the commerce of firearms with the
purpose of reducing criminality. Voting is mandatory in Brazil,
and political campaigns take advantage of free radio/TV
propaganda during the 40 days before an election. Two
political alliances arouse in the Brazilian Congress to run the
campaigns for the Yes (for the prohibition of firearm
commerce) and No (against the prohibition of firearm
commerce) votes. The media campaign stressed the benefits
and costs of each decision, trying to oppose the benefits of one
decision against the risks of the opposite one.
The pro-YES media campaign stressed, for example, that
prohibiting fire arms commerce reduces passionate murders
of beloved ones during family, neighbors, traffic, etc. quarrels,
that would result into a reduction of personal risks or costs. In
contrast, the pro-NO media campaign stressed that without
having a gun you are unable to protect yourself against
criminals, which would result into a reduction of personal
benefits. Some others media advertisement stressed either
the possible social costs of facilitating gun acquisition by
criminals by means of a free fire arms commerce or the social
benefits of an unarmed society.
Rocha et al. (2009) proposed a neuroeconomic decision-
making model, in which the adequacy or intention to
implement an action (e.g., voting) depends on the evaluation
of the risks and benefits in two different decision spaces: the
Personal (PDS) and the Interpersonal (IDS) decision spaces.
PDS is in charge of calculating the personal risks and benefits
associated with the action, and IDS is in charge of calculating
the social risks and benefits associates with a given action. In
the present case, the risks and benefits of the YES arguments
may be supposed to be evaluated in the IDS space, whereas the
risks and benefits of the NO arguments can be evaluated in the
PDS space, and then these values are used for the calculations
of the voting intention.
The time constraints imposed by the necessity of studying
a reasonable number of voters (in the present case: 32) during
a very short time (in the present case: 5 days) render EEG as the
only tool of choice for recording and analyzing the brain
activity associated with the voting decision. Because we were
not allowed to control the most important variables influenc-
ing the voting decision, we have to hypothesize that this
decision was dependent on the neural circuits involved
with self and social cognition, if the media was successful in
influencing voters.
In the analysis of event-related potentials (ERP), EEG power
spectrum and coherence (or de-coherence) are the most
frequent and widespread techniques used in EEG studies
(e.g., Alegre et al., 2006; Cohen et al., 2007; Hauk et al., 2006;
Jacobs et al., 2006; Olofsson et al., 2008; Polezzi et al., 2008).
Different ERP waveforms revealed specific amplitude and
latency characteristics related with stimulus perception:
syntactic and semantic analysis, sleep and walking states,
decision-making, etc. Different band frequency patterns are
associated with sleep and waking states, reasoning processes,
199
decision-making stages, etc. Synchronous oscillations have
been observed and investigated extensively in EEG, and
different coherence (de-coherence) indices were proposed to
quantify the synchronization between the EEG signals
recorded by different electrodes. Most of the EEG literature
on decision-making is focused on the ERP technique (Cohen
et al., 2007; Esposito et al., 2009; Heldmann et al., 2005; Polezzi
et al., 2008, 2010; Utku et al., 2002).
EEG studies of decision-making (e.g., Chen et al., 2009;
Polezzi et al., 2008a,b, 2010; Utku et al., 2002) in human
subjects have mainly focused on the:
(a) feedback-related negativity (FRN), which reflects the
medial frontal/anterior cingulate activity, and its am-
plitude codes the ongoing evaluation of events and the
prediction of future events in terms of favorable or
unfavorable outcomes;
(b) P300, which has also been reported to reflect decision-
making, and its amplitude is thought to reflect the
outcome of stimulus evaluation and decision-making.
The P300 amplitude varies with variables such as the
event probability, the motivational significance of stimuli
and the magnitude of feedback outcome, regardless of
whether the outcome is positive or negative; and
(c) N500, which is generally larger for unpredictable out-
comes, and this is thought to be generated by the posterior
cingulate cortex and the visual association cortex.
Given the necessity of scanning a large number of
volunteers in a short period of time, we selected the EEG as
the most appropriate way to record the brain activity
associated with voting decision-making. In addition, due to
the uncontrolled condition of our experiment, we had to focus
our analysis backwardly; that is, we focused on the brain
activity preceding the decision-making, whereas most, if not
all, of the experiments involving EEG and decision-making
focus on the analysis of brain activity following the decision-
making or stimulus presentation (e.g., Chen et al., 2009; Polezzi
et al., 2008a,b, 2010; Utku et al., 2002). This motivated us to use
the technique we developed (Rocha et al., 2004, 2005) before
investigating voting decision-making.
The purpose of this paper is to investigate the brain activity
associated with the voting intention declared 1 week before
the election day of the Brazilian Firearms Control Referendum,
supposing that that voting was influenced by media and
that this influence can be deciphered by neural correlates
associated with different decision spaces (PDS and IDS), and
assuming for this purpose the EEG as the tool of choice for
recording and analyzing the brain activity associated with the
voting decision. The ERP technique is used to characterize the
brain activity preceding the voting decision-making, and the
brain imaging technique developed by Rocha et al. (2004, 2005)
is applied to study the topology of the voting decision-making
brain network.
2. Results
From the total of 32 individuals studied, 10 (31%) declared the
intention to vote YES on Election Day, whereas 15 (47%)
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200 BR A I N R ES E A RC H 1 3 5 1 ( 2 01 0 ) 1 9 8 –21 1

individuals declared the intention to vote NO. The remaining Table 1 – F and p values for ANOVA mappings.
seven (22%) individuals were undecided at the time of the A ðei ; tÞ h(ei)
experiment. These voting percentages were similar to the
poll results published in the same week by Brazilian pollsters F p F p
(Ibope, 2005). Volunteers took an average of 12 s to reach a C3 1.20 0.27 37.00 0.00
decision. C4 6.05 0.01 121.00 0.00
Data analysis consisted of the following steps: CZ 95.00 0.00 121.00 0.00
F3 98.00 0.00 5.80 0.01
F4 4.30 0.02 12.11 0.00
(1) possible statistical differences for the average ampli-
F7 7.31 0.06 4.41 0.03
tude A ðei ; tÞ and the calculated entropyh(ei) concerning F8 0.00 0.82 1.51 0.02
Yes and No voters were first evaluated by means of FP1 0.00 0.82 1.93 0.21
ANOVA. The results of these analyses are shown by the FP2 97.00 0.00 2.54 0.12
ANOVA mappings in Figs. 1 and 3, plotting the statistical FZ 90.00 0.00 2.43 0.08
significance for the differences between A ðei ; tÞ of h(ei) O1 70.00 0.00 12.10 0.00
OZ 0.21 0.66 4.31 0.00
fro each electrode taking into account the type of vote as
O2 4.30 0.02 11.50 0.00
the grouping variables;
P3 0.65 0.43 11.50 0.00
(2) A ðei ; tÞ data were visually inspected (Fig. 1) for possible P4 0.32 0.52 62.00 0.00
easily identifiable components occurring at defined PZ 0.13 0.71 14.00 0.00
times before the voting decision; T3 1.36 0.25 62.00 0.00
(3) factor analysis was carried out for both A ðei ; tÞ and h(ei) T4 0.69 0.41 3.30 0.00
to disclose possible data covariation and to identify T5 29.90 0.00 15.00 0.00
T6 4.30 0.03 12.90 0.00
possible brain activity patterns associated with Yes or
No voting decisions. The results are presented in Table 1
for A ðei ; tÞ and Table 2 for h(ei). The loading factors were
used to generate the Factor Mappings shown in Fig. 2 for assumed to be different if R → 0 or inversely correlated if
A ðei ; tÞ and Fig. 3 forh(ei); R → − 1. The results showed that R → 1 or R → 0 for all
(4) linear regression analysis was used to study the possible compared mappings.
correlation between vote and h(ei), because for each
decision a unique value of h(ei) exists for each recording Fig. 1 shows the temporal evolution of A ðei ; tÞ during the
electrode. The results of this analysis are shown in two seconds that preceded the voting decision by the
Table 3, and the corresponding Regression Mapping is individuals of Class 1 or Yes voters and Class 2 or No voters.
shown in Fig. 3; and The visual inspection of Fig. 1 shows a difference for A ðei ; tÞ
(5) the Pearson's correlation coefficient Rwas calculated for evolution associated with the different voting decision,
comparing different brain mappings, such that two which is confirmed by the ANOVA shown in the inserted
mappings are expected to be similar if R → 1; they are brain mapping. This analysis showed statistically significant

Fig. 1 – Temporal evolution of A (ei ; t) during the 2 s that preceded the voting decision by the individuals of Class 1 or Yes voters
(n = 10) and Class 2 or No voters (n = 15). The inserted brain mapping color encodes the ANOVA analysis (Table 1), such that
statistically significant differences (p(ei) → 0) are depicted in bluish colors and non-significant differences (p(ei) → 1) are depicted
in reddish colors.
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BR A I N R ES E A RC H 1 3 5 1 ( 2 01 0 ) 1 9 8 – 2 11 201

–
Table 2 – Factor Analysis for A(ei,t).
Vote NO (Class 2) Vote YES (Class 1)

Extraction: principal components Extraction: principal components

hna hsim

Factor Factor Factor Factor Factor Factor

1 2 3 1 2 3

C3 0.716393 −0.03283 0.383835 C3 0.54006 0.443392 0.296677

C4 0.176105 0.687686 0.026208 C4 − 0.38913 0.224257 0.755081
CZ 0.821405 0.363859 −0.04678 CZ 0.864032 0.15836 −0.04897
F3 0.246867 0.743744 0.345671 F3 0.495682 − 0.1313 0.542722
F4 0.933855 0.122746 0.141891 F4 − 0.0771 0.887777 0.092686
F7 0.074973 0.7737 0.078913 F7 0.291887 0.706532 0.124067
F8 0.183568 −0.20251 0.544943 F8 0.213645 0.855867 −0.04571
FP1 0.024346 0.414883 0.694659 FP1 − 0.13581 0.821801 0.122069
FP2 0.146386 0.014244 0.763065 FP2 0.22034 0.701512 −0.00923
FZ 0.314326 −0.11137 −0.22925 FZ − 0.07134 0.494585 0.469489
O1 0.526566 0.528918 −0.01984 O1 0.616206 0.249811 −0.00746
OZ 0.776535 0.373813 −0.05236 OZ 0.435752 0.16587 0.555836
O2 0.580343 0.553088 −0.26819 O2 0.790489 0.184794 −0.30451
P3 0.335255 0.480346 0.03468 P3 0.878269 − 0.0293 0.282258
P4 0.849659 0.090607 0.117452 P4 0.080344 0.520613 0.542198
PZ 0.48805 0.408624 −0.12976 PZ 0.511573 − 0.09745 0.379866
T3 0.705332 0.38674 0.317334 T3 − 0.13513 0.649345 0.144688
T4 0.936426 0.072963 0.156005 T4 0.727886 0.147857 0.032165
T5 0.469806 0.514611 −0.25512 T5 0.471078 − 0.18983 0.388343
T6 0.853142 0.074734 −0.26408 T6 0.845413 − 0.18492 −0.12509
Expl.Var 6.92465 3.555576 2.07715 Expl.Var 5.362906 4.640337 2.324696
Prp.Totl 0.346232 0.177779 0.103858 Prp.Totl 0.268145 0.232017 0.116235
0.936426 0.7737 0.763065 0.878269 0.887777 0.755081
0.024346 −0.20251 −0.26819 − 0.38913 − 0.18983 −0.30451

Extraction: principal components Extraction: principal components

% total Cumul. Cumul. % total Cumul. Cumul.

Eigenvalue Variance Eigenval. % Eigenvalue Variance Eigenval. %

1 8.44703 42.23515 8.44703 42.23515 1 6.064215 30.32108 6.064215 30.32108

2 2.107497 10.53748 10.55453 52.77264 2 4.348519 21.7426 10.41273 52.06367
3 2.002848 10.01424 12.55738 62.78688 3 1.915205 9.576024 12.32794 61.6397

differences for all electrodes except FP1, F8, C3, P3, P4, PZ and
O2. It may be considered that A ðei ; tÞ, calculated for the
electrodes FP2, F3, CZ, C4 and T5 in Class 1 and for the
electrodes F4, C3, CZ, FZ, F4, C3, CZ, T3 and OZ in the case of
Class 2, exhibits a variation higher than for the remaining
electrodes during the 2-s study period. So, a distinction
between the EEG activities recorded from the individuals in
Class 1 and Class 2 can be established based on A ðei ; tÞ
calculated for the above-cited electrodes.
In contrast to the usual outcome, the present A ðei ; tÞ analysis
did not disclose any conspicuous negatively or positively isolated
components that could be easily identified with a defined latency
either for Class 1 (Yes vote) or Class 2 (No vote). Because of
this lack of specific waveform with a distinctive (negative)
latency identified for either the YES on the NO decision-making,
we decided to use Principal Component Analysis to understand
the possible differences of brain activity associated with these
distinct decision-making directions (Fig. 2).
Fig. 2 shows the brain mappings depicting the results of
A ðei ; tÞ Factorial Analysis for Classes 1 and 2, shown in Table 1.
Three factors (Fi) explained around 60% of the total variance
of A ðei ; tÞ in Class 1 and Class 2, respectively (Table 2). The
eigenvalues for these factors varied from 1.91 for F3 of Class 1
to 8.44 for F1 of Class 2, stressing the robustness of the
analysis.
The Pearson's correlation coefficient between these
factors calculated for Classes 1 and 2 revealed the existence
of three patterns of A ðei ; tÞ covariation (Patterns Pi in Fig. 2).
Each pattern P i is defined by the pair of factors F i ,
Fjhaving the highest Ri, j values (R = 0.63, 0.34 and 0.21 in
Fig. 2). Pattern P1 is composed by factors F3 from Class 1
(eigenvalue of 1.91) and F2 from Class 2 (eigenvalue of 2.10),
whose similarity is at the level of R = 0.63. Pattern P1 has C4
as the common high-loading electrode (loading factor
greater than 0.6) for both Classes 1 and 2, with F3 and F7
for Class 2 only. Pattern P2 is composed by factors F1 from
both Class 1 (eigenvalue of 6.06) and Class 2 (eigenvalue of
8.44), whose similarity is at the level of R = 0.34. Pattern P2
has CZ, T4 and T6 as the common high-loading electrodes
for both Classes 1 and 2, has T4, P3, O1 and O2 as high
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202 BR A I N R ES E A RC H 1 3 5 1 ( 2 01 0 ) 1 9 8 –21 1

Fig. 2 – A (ei ; t) Factorial brain mappings for Class 1 (Yes vote) and Class 2 (No vote). The normalized values f j (ei ) of the factor
loadings fj(ei) (Table 1) for each electrode (ei) of the three principal components Fj (eigenvalues greater than 1) are color-coded
such that f j (ei )→1 is colored blue and f j (ei )→0 is colored red. R-values represent the Pearson's correlation coefficient between
each pair of electrodes.

loading electrodes for Class 1 and has F4, C3, T3, P4 and OZ
for Class 2. Pattern P3 is composed by factors F2 from Class 1
(eigenvalue of 4.34) and F3 from Class 2 (eigenvalue of 2.00),
whose similarity is at the level of R = 0.21. Pattern P3 has FP1
and FP2 as the common high-loading electrodes for both
Classes 1, and F4, F7 and F8 are the high-loading electrodes
for Class 1.
Fig. 3 shows the brain mappings depicting the results of
h (ei) factorial and ANOVA analyses for Classes 1 and 2,
shown in Table 2. The values of h(ei)calculated for Class 1
and Class 2 were different for all electrodes except Fp1, Fp2
and FZ. Three factors explained around 80% of the total h(ei)
variance in Class 1 and Class 2, respectively (Table 2). The
eigenvalues for these factors varied from 1.30 for F3 of Class
1 to 12.08 for F1 of Class 1, stressing the robustness of the
analysis.
The Pearson's correlation coefficient between these
factors calculated for Classes 1 and 2 revealed the existence
of three patterns of h(ei) covariation (Patterns Pi in Figure 3).
Each pattern Pi is defined by the pair of factors Fi, Fj having
the highest Ri, j values (R = 0.68, 0.57 and 0.03 in Fig. 3).
Pattern P1 is composed by factorsF3 from Class 1 (eigenval-
ue of 1.3) and F1 from Class 2 (eigenvalue of 10.1), whose
similarity is at the level of R = 0.68. Pattern P1 has T3, T4, T5
and T6 as the common high-loading electrodes (loading
factor greater than 0.6) for both Classes 1 and 2, and it was
P4 for Class 1 or F3, O1, O2 and OZ for Class 2. Pattern P2 is
composed by factors F2 from Class 1 (eigenvalue of 3.4) and
F3 from Class 2 (eigenvalue of 3.2), whose similarity is at
the level of R = 0.57. Pattern P2 has FP1 and FP2 as the
common high-loading electrodes for both Classes 1 and 2,
and F4 and F8 are the high-loading electrodes for Class 1
only. Pattern P3 is composed by factors F1 from Class 1
(eigenvalue of 12.0) and F2 from Class 2 (eigenvalue of 3.5),
whose similarity is at the level of R = 0.03. Pattern P3 has C3,
C4, Cz and P3 as the common high-loading electrodes for
both Classes 1 and 2, and F3, FZ, PZ, O1, O2 and OZ are the
high-loading electrodes for Class 1 and F4, F8, T6 and P4 are
for Class 2.
Fig. 3 also shows the brain mappings depicting the results
of the h(ei)regression analysis for Classes 1 and 2, shown in
Table 3. The values of R2 for these regressions were 0.57 and
0.51, respectively. Their statistical significances were p = 0.09
and p = 0.01, respectively. The decision of voting Probably NO
(green) and Certainly NO (blue) increased as h(ei) augmented
at the electrodes Fp2, F4 and F8. The decision of Certainly
Not voting NO (red) increased as h(ei) augmented for
electrodes Fp1, F3, F7, T4, P3, P4, O2 and OZ. The decision
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BR A I N R ES E A RC H 1 3 5 1 ( 2 01 0 ) 1 9 8 – 2 11 203

Fig. 3 – h(ei)regression brain mappings (Table 4) on the left, factorial brain mappings (Table 3) for Class 1 (Yes vote) and Class 2
(No vote) in the middle and ANOVA brain mapping on the right (Table 1). The normalized values β (ei ) of the angular coefficients,
BETA, for each electrode (ei) are color-coded such that β (ei )→1 is colored blue and β (ei )→0 is colored red. According to the
normalization rule, β(ei )>0:5 implies BETA > 0 and β (ei )<0:5 implies BETA < 0. The normalized values f j (ei ) of the factor loadings
fj(ei) (Table 2) for each electrode (ei) of the three principal components Fj (eigenvalues greater than 1) are color-coded such that
f j (ei )→1 is colored blue and f j (ei )→0 is colored red. The R-values represent the Pearson's correlation coefficient calculated
between each pair of electrodes. The p(ei) values of the ANOVA brain mapping are color-coded such that p(ei) → 0 is colored blue
while p(ei) → 1 is colored red. Brain contours are simply used as references for the topographical location of the 10/20 system
electrodes. CN in the middle bar means Certainly NO Vote, and NN means Certainly Not NO Vote. CY means Certainly YES
Vote, and NY means Certainly Not YES Vote.

of voting Probably YES (green) and Certainly YES (blue)
increased as h(ei) augmented at the electrodes Fp2, F3, F4, FZ,
C3, CZ, PZ, T6 and OZ. The decision of Certainly Not voting
YES (red) increased as h(ei) augmented for electrodes P3, T5
and O1.
The regression analysis for h(ei)Pi and A ðei ; tÞPi showed that:
tionship is found when we regress A ðei ; tÞPi as a function of
h(ei)Pi.
Finally, logistic regression between YES(=1)/NO(=0) vote
and h(ei)Pi and A ðei ; tÞPi explained 40% and 60% of their
variance, respectively.

hðei ÞP1 = 0:29 + 0:16A ðei ; tÞP1 + 0:34A ðei ; tÞP2 −0:07A ðei ; tÞP3 ;
r = 0:35 ðp < 0:17Þ; 3. Discussion

hðei ÞP2 = 0:87 + 0:05A ðei ; tÞP1 −0:57A ðei ; tÞP2 −0:17A ðei ; tÞP3 ;
To the best of our knowledge, this is the first study to
r = 0:55 ðp < 0:003Þ;
investigate the neural process supporting a real, complex
hðei ÞP3 = 0:18−0:15A ðei ; tÞP1 + 0:40A ðei ; tÞP2 −0:38A ðei ; tÞP3 ; and non-controlled decision-making situation such as voting
r = 0:44 ðp < 0:04Þ decision-making. Most of the recently published studies about
complex decision-making deal with experiments in well-
Note that there is a linear relationship between h(ei)Pi controlled laboratories (e.g., Esposito et al., 2009; Goldman
and A ðei ; tÞPi. In contrast, no reasonably good linear rela- et al., 2009; Ino et al., 2010) or simulated hypothetical conditions
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204 BR A I N R ES E A RC H 1 3 5 1 ( 2 01 0 ) 1 9 8 –21 1

Table 3 – Factor Analysis for h(ei).

Vote NO (Class 2) Vote YES (Class 1)

Factor loadings (Varimax raw) Factor loadings (Varimax raw)

Extraction: principal components Extraction: principal components

Factor Factor Factor Factor Factor Factor

1 2 3 1 2 3

C3 0.303021 0.756425 0.328989 C3 0.714671 0.05566 0.573831

C4 0.169343 0.953117 0.179718 C4 0.859526 0.017209 0.270431
CZ 0.394629 0.882466 0.030144 CZ 0.924638 0.08137 0.008911
F3 0.702341 0.531177 0.27285 F3 0.841299 0.267794 −0.14193
F4 0.365894 0.831675 0.237033 F4 0.254855 0.788703 0.092138
F7 0.395954 0.519182 0.430755 F7 0.024754 0.561752 0.41611
F8 −0.05468 0.778331 0.513248 F8 − 0.01893 0.867088 −0.04026
FP1 0.319527 0.354102 0.552115 FP1 − 0.09042 0.890918 0.108808
FP2 0.074725 0.450829 0.623862 FP2 0.061982 0.940355 −0.0032
FZ 0.113395 0.156293 0.83417 FZ 0.707317 0.419191 0.208262
O1 0.930001 0.231891 0.183899 O1 0.836588 −0.03039 0.438318
O2 0.870292 0.377368 0.253838 O2 0.901654 −0.03967 0.289379
OZ 0.790207 0.561794 0.071003 OZ 0.802363 0.109763 0.12503
P3 0.185154 0.901274 0.114804 P3 0.871922 −0.0704 0.3183
P4 0.583509 0.743501 −0.13627 P4 0.114162 0.153884 0.844156
PZ 0.859247 0.117221 −0.03408 PZ 0.734084 −0.08941 0.579657
T3 0.914068 0.265075 −0.05751 T3 0.08191 0.085199 0.931874
T4 0.976257 − 0.01251 0.054623 T4 0.56197 0.143208 0.73699
T5 0.92801 0.237388 0.019148 T5 0.509427 −0.02971 0.771146
T6 0.921141 0.217318 0.17597 T6 0.656472 −0.15516 0.604961
Expl.Var 8.041811 6.526223 2.284749 Expl.Var 7.872314 3.728809 4.47472
Prp.Totl 0.402091 0.326311 0.114237 Prp.Totl 0.393616 0.18644 0.223736
0.976257 0.953117 0.83417 0.924638 0.940355 0.931874
−0.05468 − 0.01251 −0.13627 − 0.09042 −0.15516 −0.14193

No Yes

Extraction: principal components Extraction: principal components

% total Cumul. Cumul. % total Cumul. Cumul.

Eigenvalue Variance Eigenval. % Eigenvalue Variance Eigenval. %

1 10.17027 50.85135 10.17027 50.85135 1 12.08706 60.43529 12.08706 60.43529

2 3.548731 17.74366 13.719 68.59501 2 3.456356 17.28178 15.54341 77.71707
3 2.224301 11.12151 15.9433 79.71651 3 1.30937 6.54685 16.85278 84.26392

(e.g., Robertson et al., 2007; Kliemann et al., 2008; Young and
Saxe, 2008).
Given the necessity of scanning a large number of volunteers
in a short period of time, we selected the EEG as the most
appropriate way to record the brain activity associated with the
voting decision-making. In addition, due to the uncontrolled
condition of our experiment, we had to focus our analysis
backwardly; that is, we focused on the brain activity preceding
the decision-making, whereas most, if not all, of the experi-
ments involving EEG and decision-making focus on the analysis
of the brain activity following the decision-making or stimulus
presentation (e.g., Utku et al., 2002; Cohen et al., 2007; Polezzi et
al., 2008a,b). This motivated us to look for new methods for EEG
analysis that could be informative without being dependent on
the recognition of defined wave forms with characteristic
latency, as usually done in ERP analysis.
It is possible to conclude, from the new ERP analysis
proposed here, that the vote decision involved a set of widely
distributed neurons as a part of a complex processing network in
charge of calculating benefits and risks of the decision of
prohibiting or allowing firearm commerce. The possible benefits
and risks of each decision were the focus of the media campaign
in radio, TV and newspapers. As mentioned in Introduction, the
pro-YES media campaign stressed, for example, that prohibiting
fire arms commerce reduces passionate murders of beloved
ones during family, neighbors, traffic, etc. quarrels, that would
result into a reduction of personal risks or costs. In contrast, the
pro-NO media campaign stressed that without having a gun you
are unable to protect yourself against criminals, which would
result into a reduction of personal benefits. Some others media
advertisement stressed either the possible social costs of
facilitating gun acquisition by criminals by means of a free fire
arms commerce or the social benefits of an unarmed society.
The voters were then asked to consider the benefits for one (YES
or NO) decision and the risk of the opposite (NO or YES) decision
in order to decide their vote. So, it may be hypothesized here that
our volunteers were influenced by the media campaign, and
their voting decision was based upon the comparison of the
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BR A I N R ES E A RC H 1 3 5 1 ( 2 01 0 ) 1 9 8 – 2 11 205

20
Table 4 – Regression analysis v=τ+∑ βihv(Rj).
i=1

Regression summary for dependent variable: v

R = 0.47401843, R2 = 0.22469347, Adjusted R2 = 0.21720620

F(20.2071) = 30.010, p < 0.0000, Std. Error of estimate: 1.6052

St. Err. St. Err.

BETA of BETA B of B t(2071) p-level h ðei Þ h ðei Þbi

Intercept 73.4995 0.092447 795.0415 0

C3 0.213719 0.044681 0.268551 0.056144 4.783253 1.85E-06 2.048764 0.550198

C4 − 0.18256 0.040078 − 0.21511 0.047223 −4.55522 5.54E-06 2.320727 -0.49921
CZ − 0.1403 0.044575 − 0.17471 0.055509 −3.14745 0.00167 2.217982 -0.3875
F3 − 0.35163 0.034037 − 0.45556 0.044098 − 10.3307 1.98E-24 2.3037 -1.04947
F4 0.070889 0.035143 0.100321 0.049734 2.017142 0.043809 2.344191 0.235173
F7 − 0.06023 0.02748 − 0.10254 0.046781 −2.19192 0.028496 1.698536 -0.17417
F8 0.128032 0.031324 0.20291 0.049643 4.08735 4.53E-05 1.832645 0.371862
FP1 0.33265 0.034048 0.479484 0.049077 9.769934 4.51E-22 2.092309 1.003228
FP2 − 0.31124 0.03965 − 0.40437 0.051514 −7.84967 6.63E-15 2.250064 -0.90985
FZ 0.034281 0.032449 0.038471 0.036415 1.056467 0.290878 2.498155 0.096107
O1 − 0.72118 0.172827 − 0.22476 0.053864 −4.17281 3.13E-05 2.143427 -0.48176
O2 1.063042 0.175132 0.331745 0.054654 6.069933 1.52E-09 2.197273 0.728935
OZ − 0.25695 0.147649 − 0.07995 0.045941 −1.74026 0.081962 1.963082 -0.15695
P3 − 0.0072 0.036996 − 0.01012 0.052025 −0.1946 0.845726 1.876827 -0.019
P4 − 0.07842 0.033514 − 0.11801 0.050434 −2.33995 0.019381 1.742227 -0.2056
PZ − 0.01659 0.03997 − 0.02012 0.048492 −0.41496 0.678213 1.819464 -0.03661
T3 0.138843 0.033612 0.225105 0.054496 4.130688 3.76E-05 1.671445 0.376251
T4 0.156891 0.03378 0.227492 0.048981 4.644515 3.62E-06 1.570627 0.357305
T5 0.037126 0.031826 0.065031 0.055748 1.166522 0.243538 1.802555 0.117223
T6 − 0.173 0.053258 − 0.05039 0.015512 −3.24834 0.001179 1.772145 -0.08929

calculated benefits and risks to calculate the adequacy of each
voting alternative. If this is true, then the differences of A ðei ; tÞ
covariance observed between YES and NO Voting for patterns P2
and P3 could reflect the differences in the corresponding
calculated benefits and risks of each decision. In addition, it
may be also hypothesized that voting decision-making required
recursive calculations of the benefits and risks for different
arguments in favor of both voting alternatives, such that no
specific waveform with a distinctive (negative) latency is easily
identified either for the YES on the NO decision-making.
The Principal Component Analysis also revealed the exis-
tence of three different patterns (Pi) (Fig. 3) of brain activity
concerning h(ei) that explain 80% of the data covariance
associated with the vote decision. If it is considered that the
different patterns Pi reflect the structure of the brain networks
involved in the reasoning supporting the voting decision, then it
may be concluded from the Principal Component Analysis that
the YES and NO voting reasoning differences are mostly
supported by the differences in network organization charac-
terized by patternP3. This hypothesis seems to be further
substantiated by the linear multiple regression analysis corre-
lating the vote decision v and h(ei), showing that voting YES or
NO was associated with high values of h(ei) calculated for distinct
set of electrodes. Therefore, it may be proposed here that: (a) the
distinct networks disclosed by the distinct NO and YES voting h
(ei) pattern P3 would support the distinct arguments for and
against each voting decision; and (b) the similar networks
disclosed by similar h(ei) patterns P1 and P2 would be enrolled in
the calculations of the risks and benefits associated with the
selected arguments and the adequacy of each voting decision.
At this point, it would be convenient to propose a model
for the voting decision-making in the case of a referendum.
First, however, it should be mentioned that a referendum is
a particular case of election in which only two alternatives
about a single problem are presented to potential voters.
Therefore, this is a rather simple type of voting decision, and it
can be done by contrasting two sets of arguments supporting
or rejecting each voting option. This was the case in the
Brazilian firearm referendum. Two political coalitions were
formed, and two sets of arguments were elaborated and
discussed. The media campaign worked out this process
intensely asking the voters to evaluate the personal/social
costs/benefits of each possible voting decision to calculate the
adequacy of each of these decisions.
In line with these assumptions, the decision model intro-
duced by Rocha et al. (2009) proposes the adequacy or intention
to implement an action (e.g. voting) depends on the evaluation
of risks and benefits in two different decision spaces: the
Personal (PDS) and the Interpersonal (IDS) decision spaces. The
adequacy (or fairness) of the action (voting YES or NO) was
proposed to be dependent on the balance between the ratios
personal risk/social benefit and the social risk/personal benefits.
The adequate (fair) action is supposed to provide high social and
personal benefits at low personal and social risks. Whenever the
ratio risk/benefit approaches one, conflict about acting or not
arises and makes the decision-making difficult. Conflict is a
source for procrastination, that is, in the case of voting, is the
source for undecided and certainly not votings. Therefore, the
willingness to act (intention to vote) depends on adequacy and
conflict. In the present case, the risks and benefits of the YES
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206 BR A I N R ES E A RC H 1 3 5 1 ( 2 01 0 ) 1 9 8 –21 1
arguments are evaluated in the IDS space, whereas the risks and
benefits of the NO arguments are evaluated in the PDS space,
and then these values are used for the calculations of the voting
intention.
In this context, we may propose that the networks
identified by the different h(ei)P3 patterns are those supporting
the risk and benefits evaluation in PDS and IDS, whereas the
similar networks disclosed by h(ei)P1 and h(ei)P2 patterns are in
charge of calculating the adequacy or intention of each voting
decision. Being more specific, we may propose that the pattern
h(ei)P2 is associated with the executive network in charge of
controlling the calculations, whereas pattern h(ei)P1 is associ-
ated with this calculation.
Dosenbach et al. (2006) and Dosenbach (2007) proposed that
mixed blocked/event-related fMRI allow the direct and separate
extraction of three types of task set-related signals: (1) a signal
tied to the start of a specific task condition, which should be, at
least in part, related to the instantiation of task parameters;
(2) activities sustained at a constant level across the task period,
some of which likely reflect task set maintenance; and (3) an
error-related feedback, to monitor processing and encourage
adjustments of top-down signals. Using data from ten different
experiments and 183 subjects, these authors (Dosenbach et al.,
2008) confirmed their propositions and identified, besides a set
of task-related signals, the presence of two different networks
related with task control and control maintenance: the fronto-
parietal and the cingulo-opercular networks. The fronto-parietal
network includes the dorsolateral prefrontal cortex and intra-
parietal sulcus, and it seems to initiate and adjust the control of
the task-solving cognitive mechanism. The cingulo-opercular
net includes the dorsal anterior cingulated/medial superior
frontal cortex, anterior insula/frontal operculum and anterior
prefrontal cortex, and it provides the stability for this mecha-
nism for the duration of the task. In addition, another network
would provide the instantiation of task parameters used by the
fronto-parietal network. In addition, Fair et al. (2009) showed
that brain processing changes from a more local to a more
distributed one from early school-age children to adults, while
maintaining “small-world”-like properties. Here, we propose
that h(ei)P1 and h(ei)P2 are the EEG signatures of the fronto-
parietal and cingulo-opercular networks, and h(ei)P3 is associat-
ed with the instantiating network setting the initial parameters
for calculating task-solving.
Recent studies involving EEG spectra analysis has provided
support for identifying different networks associated with the
most common band frequencies (Chen et al., 2008; Ferri et al.,
2008; Mantini et al., 2007) and have shown that each fMRI-
identified network is characterized by a specific electrophys-
iological signature that involves the combination of different
brain rhythms. Scheering et al. (2008) correlated frontal theta
activity with default network activity. Chen et al. (2008)
showed that the spectral EEG default network is characterized
by a complex EEG spectral field dominance, with delta, theta,
beta-2 and gamma activities predominating at the frontal
electrodes, while alpha1, alpha2 and beta-1 activities pre-
dominated at the posterior electrodes. A complex but specific
correlation between resting state networks and the EEG
default network is reported by Mantini et al. (2007). The
relationships between h(ei)Pi and A ðei ; tÞPi disclosed by regres-
sion analysis adds new information to the understanding of
the relations between the EEG activity and the brain network
organization. However, further experiments involving, for
instance, simultaneous EEG and fMRI recordings are necessary
to provide a better support for this hypothesis.
If our hypothesis holds, then h(ei)P3 differences for Yes and
No voters shall reflect the choices of the different arguments
for supporting their distinct voting decisions. Taking into
consideration that the media campaign proposed YES argu-
ments to be evaluated in the IDS space, whereas the risks and
benefits of the NO arguments should be evaluated in the PDS,
then the Yes h(ei)P3 is associated with the neural circuits
composing the IDS space and the No h(ei)P3 is associated to
those composing the PDS space.
Kennan (2000) reviewed the literature on fMRI, ERPs,
transcranial stimulation, split-brain and focal lesions and
proposed that self-recognition, a PDS-specific task, shows that
the right prefrontal cortex may be an important component of a
neural circuit for self-evaluation. Udin (2004) showed that
images containing more “self” information increased activity
in the right hemispheres, including the inferior parietal lobe,
inferior frontal gyrus and inferior occipital gyrus, whereas
images containing more “other” information increased signals
in the medial prefrontal cortex and precuneus. Iacoboni (2004)
studied people watching realistic movie clips depicting everyday
social interactions, and they observed that the movie clips'
epochs showing people interaction, as compared to those
depicting single individuals involved in daily activities, yielded
increased activity in the medial parietal (precuneus) and
dorsomedial prefrontal cortices. Those authors proposed that
default networks would participate in social cognition. A similar
proposition was also made by Schilbahc (2008). The ability to
differentiate kin from non-kin members of one's social group is
paramount to survival when living in social groups. Platek and
Kemp (2009) showed that kin recognition is associated with the
posterior cingulate cortex and cuneus. All of these data are in
accordance with our results showing that No h(ei)P3 is charac-
terized mostly by high h(ei)values for the right anterior central
and frontal electrodes, whereas Yes h(ei)P3 depicts high h(ei)
values mostly for the medial electrodes.
From what was discussed so far, we may assume that:
(1) h(ei)P3 is associated with the networks evaluating
benefits (b(vi)) and risks (r(vi)) at both PDS and IDS
networks;
(2) h(ei)P2 is associated with the Voting Processing Network
calculating the adequacy (ψ(vi)) of voting YES or NO
taking into account the calculations done by the PDS
and IDS networks; ; and
(3) h(ei)P2 is associated with the Voting Decision Network
determining the intention of voting YES or NO taking
into account the adequacy and conflict estimated by the
Voting Processing Network.
4. Conclusion
We claimed at the Introduction that the study of real and
natural decision-making imposes some constraints over the
experimental paradigms and technologies used to collect the
required data for analysis and decision-making modeling. We
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BR A I N R ES E A RC H 1 3 5 1 ( 2 01 0 ) 1 9 8 – 2 11 207

have stressed that such kind of study does not allow the
control of the most important variables by the researcher, that
in the majority of the cases it requires a portable recording
technique and that the experiments have to be done as close
to the actual moment of the decision-making as possible, if
not at the very moment. If the last is not the case, we have to
study the intention or willingness of deciding instead of the
real decision-making.
Here, we studied the voting intention (or willingness) in the
Brazilian Firearm Commerce Referendum during the week
preceding the election, using two different EEG methodologies
(Event Related Activity and Event Related Entropy), and we
correlated the results with the media campaign, which linked
voting decision with the evaluation of the cost and benefits in
both personal and social contexts.
Because we were not allowed to control the most important
variables influencing the voting decision, we had to hypoth-
esize that this decision was to be dependent on the neural
circuits involved with self and social cognition, if the media
was successful in influencing voters.
Our results show that voting decision-making involves a
very complex network of widely distributed neurons in charge
of: (1) evaluating the risk and benefits in both Personal and
Interpersonal Decision Spaces; (2) calculating the fairness
of each voting decision; and (3) determining the intention
(or willingness) to vote Yes or No or not to vote. The results
validate, therefore, the EEG technologies used here as the tool
of choice for such a study type, and they show that if the
important variables influencing the decision-making cannot
be controlled by the researcher, they can be learned from the
environmental conditions under which the decision is sup-
posed to be made. This kind of approach paves the way for
future studies of human decision-making for the complexes
issues of a modern society.
5. Experimental procedures
Thirty-two subjects, 16 female and 16 male adults (Table 5),
had their EEG registered with 20 electrodes placed according to
the 10/20 system; impedance was smaller than 10 Kohm; the
low band pass filter was at 50 Hz; the sampling rate was
256 Hz; and there were 10 bits of resolution while selecting one
of the possible options,
1) I will certainly vote YES (CY)
2) I will probably vote YES (PY)
3) I will certainly not vote YES (NY)
4) I am undecided (UN)
5) I will certainly not vote NO (NN)
6) I will probably vote NO (PN)
7) I will certainly vote NO (CN),
reporting their intended votes in the real election to be held
one week later. For the purpose of calculations, the possible
answers were numerically coded as:
CY = 1; PY = 2; NN = 3; UN = 4; NY = 5; PN = 6 and CN = 7;
and they were classified as: Class 1 or Yes vote (comprising
codes 1, 2 and 3) and Class 2 or No vote (comprising codes 5,6
and 7).
Two networked personal computers were used, one for the
EEG recording and the other for presenting the voting options.
The volunteers were allowed to take as much time as needed
to vote. The timing of voting options' presentation (to) and
voting decision (tv) were tagged as corresponding marks
(mo, mv) in the file of the recorded EEG. The EEG was visually
inspected for artifacts before being processed, and the events
associated with a bad EEG (e.g., when eye movements could
compromise the regression analysis) were discarded. The
voting decision time was calculated as tv − to and averaged 32 s.
The EEG recorded during the two seconds preceding the voting
decision was used for both the ERP analysis and the brain
mapping (Foz et al., 2001; Rocha et al., 2004, 2005).
The ERP was calculated as the average A ðei ; tÞ of the
amplitudeA(ei, t) of the EEG activity recorded for each electrode
averaged over the 2-s epoch preceding the voting decision
(therefore, t ∈ ½−2; 0Š). The correlation coefficient ri, jcomputed
for the EEG amplitude recorded by the electrodes ei and ej was
used as a measure of the connectivity w(ni, nj) to compute the
net entropy h(ei) according to Rocha et al. (Foz et al., 2001;
Rocha et al., 2004, 2005). One-way Analysis of Variance
(ANOVA) was used to compare A ðei ; tÞ and h(ei), with classifica-
tions variables of the above-mentioned Classes 1 and 2.
Factorial analysis was used to study the covariation of
A ðei ; tÞ and h(ei) for the groups of volunteers that voted Yes
(Class 1) and No (Class 2). This analysis (Tables 2 and 3)
revealed that three factors accounted for more than 60% of
covariation of A ðei ; tÞ or h(ei) for both Class 1 and Class 2. The
resulting eigenvalues for all factors were greater than 1.9.
Three factorial brain mappings (Figs. 2 and 3) for the Yes and
No voters were constructed to describe the results of these
factorial analyses. These mappings were constructed taking
into account the loading values fj(ei) of A ðei ; tÞ or h(ei) for factors
Fj( j = 1, 2, 3). To estimate possible similarities between the
loadings fj(ei)of Fj( j = 1, 2, 3) in Class 1 and Class 2, we used
Pearson's correlation coefficients.
Linear regression analysis was used to estimate the
20
function v = τ + ∑ βi hðei Þ, correlating the vote v and the net
i=1
entropy h(ei) (see Table 5). A regression brain mapping (Fig. 3)
was constructed taking into account the normalized values of
βh ðei Þ, where h ðei Þ is the average of the net entropy h(ei)
calculated for all subjects.

Table 5 – Characteristics of the studied population.

Age Gender Instruction Income Appendix 1. Graph theory
Mean 35 Female 0.42 Primary School 0.17 Low 0.41
A graph G = ðN; A; WÞ is a mathematical description of a
Max 63 Male 0.58 High School 0.30 Medium 0.44
network, consisting of a collection N of nodes n, A of arcs a
Min 18 College 0.53 High 0.27
(n i , n j )connecting the nodes n i , n j and W of weights of
 Author's personal copy
208 BR A I N R ES E A RC H 1 3 5 1 ( 2 01 0 ) 1 9 8 –21 1
connections w(ni, nj)of a(ni, nj). The average weighting connec-
tion wðni Þ of the node ni is calculated as:
c−1
∑ wðni ; nj Þ
j=1
wðni Þ =
c−1
where c is the cardinality of G.
The following are important definitions in graph theory.
A pathway p is an ordered set of nodes ni, nm for which
w(nj, nj + 1) > 0, i ≤ j < m. A closed pathway is defined for ni = nm. The
length l(p)of a pathway p is equal to the number k = m − i of arcs
a (ni, nj)connecting the nodes ni, nj. The power π(p)of a pathway p
−1
is defined by its weakest link, that is π(p) = min m j = i w(nj, nj + 1). A
pathway p is called preferential over p' if π(p) > π(p ' ).
A graph is called direct if w(ni, nj) ≠ w(nj, ni) for at least a pair
of nodes ni, nj ∈ N. A graph is called strictly direct if w(ni, nj) ≠ w
(nj, ni) for all pairs of nodes ni, nj ∈ N.
Information is allowed to flow from ni to nj if w(ni, nj) > 0. As a
consequence, a pathway is a channel for information distri-
bution. Here it is proposed that the maximum amount of
information h(ni, nj) that may be transmitted from ni to nj is
constrained by w(ni, nj) and it is calculated as:
hðni ; nj Þ = −wðni ; nj Þlog2 wðni ; nj Þ−ð1−wðni ; nj ÞÞlog2 ð1−wðni ; nj ÞÞ
The average amount h ðni Þ of information handled by ni is
proposed to be calculated as:
h ðni Þ = −w ðni Þlog2 w ðni Þ−ð1−wðni ÞÞlog2 ð1−wðni ÞÞ
The maximum amount of information h(p) that may be
transmitted through pathway p is given by:
hðpÞ = −πðpÞlog2 πðpÞ−ð1−πðpÞÞlog2 ð1−πðpÞÞ
and the total amount h(ni) of information handled by node ni is
given by:
c−1
hðni Þ = ∑ h ðni Þ−hðni ; nj Þ
j=1
The network efficiency has been expressed in terms of
information flow (Marchiori and Latora, 2000) and it depends
on the structure of the network.
The concept of “small-world” is strongly related to the
average shortest path length (L) and clustering coefficient (C)
concepts (Iturria-Medina et al., 2008). A cluster σ is a subset of
the network in which vertices adjacents to any vertex are
adjacent to each other (Wats and Strogatz, 1998). The small-
world network is characterized by a high clustering coefficient
and a small average path length. The scale free network
studied by Barabasi and Albert (1999) was the second major
step for the understanding of the growth of the real networks.
The authors proposed a model for the growth of a network
where p for newly added edges to a node (or vertex) depends
upon the degree k of this vertex, such that vertices with a large
number of edges (high k), called hub nodes, are more likely to
get even more nodes, that isp(k) = k − α. The exponent is
expected to vary in the range2 < α < 3. Broad-scale networks
are characterized by a degree distribution that has a power law
regime followed by a sharp cutoff that restricts the increase of
p(k). The cutoff function constrains the maximum number of
nodes that may connect to hub nodes (Iturria-Medina et al.,
2008). For example p(k) = k − α log(k / kc)is a broad-range network,
where kc is the limiting degree. The high connectivity of hubs
reduces the average path length of a network.
According to the cluster definition w(ni, nj) = w(nj, ni) → 1 for
all ni, nj ∈ σ. Therefore h(ni, nj) → 0. Now, if it is required that
wðni Þ≅0:5, then h ðni Þ≅1. In such a condition, h(ni) → k.
Appendix 2. Brain networks
Many functional neuroimaging studies, have frequently
shown task-induced signal attenuation compared to a control
state (e.g., Raichle et al., 2001). Among the brain areas with
decreased brain activity, the anterior (ACC) and posterior
cingulate cortex (PCC), precuneus (PCu), medial prefrontal
cortex (MPFC), left inferior temporal cortex (ITC) and bilateral
inferior parietal cortex (IPC) often exhibit decreased brain
activity from a baseline state other than from an activated
state. During rest but awake state, the areas within this default
mode network have cerebral blood and cerebral metabolism
rate for oxygen levels significantly above the global mean, but
have oxygen extraction fraction level similar to the global
mean (Raichle et al., 2001). This set of areas are proposed to
compose the so called default network. PCC/PCu and ACC/MPFC
constitute the main elements of this default mode network
(Greicius et al., 2003; Raichle et al., 2001).
Subsequently, a large set of studies has showed the existence
of a default mode network and its anti-correlation network
which showed negative linear correlation with the default mode
network (Fransson, 2005; Fox et al., 2005) during a resting-state
using functional magnetic resonance imaging (fMRI). Kelly et al.
(2008) quantified the negative correlation between these two
networks in 26 subjects, during active (Eriksen flanker task) and
resting state scans, and found that the strength of the
correlation between the two networks varies across individuals.
These individual differences appear to be behaviorally relevant,
as interindividual variation in the strength of the correlation
was significantly related to individual differences in response
time variability: the stronger the negative correlation (i.e., the
closer to 180° antiphase), the less variable the behavioral
performance. This relationship was moderately consistent
across resting and task conditions, suggesting that the measure
indexes moderately stable Individual differences in the integrity
of functional brain networks. Fransson (2006) compared intrin-
sic activity during rest and the two-back task to the signal
increases and decreases observed in an epoch-related version of
the working memory task. His results show that spontaneous
intrinsic activity in the default-mode network is not extin-
guished but rather attenuated during performance of the
working memory task. Moreover, he showed that the intrinsic
activity in the task-positive network is reorganized in response
to the working memory task. He concluded that his results
indicates that task-induced signal deactivations in the default-
mode regions is modulated by cognitive load and that intrinsic,
spontaneous signal fluctuations in the default-mode regions
persist and reorganize in response to changes in external work
load.
Graph theory allows the definition of what should be
considered an optimal network. The notion of an optimal
 Author's personal copy
BR A I N R ES E A RC H 1 3 5 1 ( 2 01 0 ) 1 9 8 – 2 11
network is closely associated with the “small-world” phenom-
enon (Reijneveld et al, 2007; Wats and Strogatz, 1998). The so-
called “small-world” network architecture is distinguished
from either ordered or random networks. On average, a
sparsely connected graph is expected to have a lower
clustering coefficient and longer path length than a densely
connected one with the same topology. Networks with ‘small-
world’ architecture are characterized by a combination of
strong local clustering and a short characteristic path length
(an index of global integration). This means that although most
of the connectivity is local, the network remains highly
integrated due to a small number of long distance connections.
Networks with “scale-free” architecture (Barabasi and Albert,
1999) are characterized by the presence of nodes with very
large number of long distance connections (the hub nodes).
The likelihood p(k) of a node to have k connections is given by p
(k) = k − α,2 < α < 5. Broad-scale networks are characterized by a
degree distribution that has a power law regime followed by a
sharp cutoff that restricts the increase of p(k). The cutoff
function constrains the maximum number of nodes that may
connect to hub nodes (Iturria et al, 2008). For example p(k) = k1
−α
log(k / kc) is a broad-range network, where kc is the limiting
degree. From equations 4, broad-range and scale-free networks
have clusters of well-connected nodes ei and ejfor which pi,
j → 0.5 because the characteristic pathway length li, j tends to 2,
and a small number of hub nodes eh for which ph, j → 1 if k < kc.
Micheloyannis et al. (2006) recorded EEG signals to study
neuronal interactions during working memory tests in indivi-
duals who had few years of formal education (LE) as compared
to individuals with university degrees (UE). They quantified the
synchronization between EEG channels in several frequency
bands, and then converted EEG signal correlations into graphs to
estimate the clustering and distance characteristics of the
underlying processing networks. According to the authors,
their finding supports the neural efficiency hypothesis and
suggests that the connections between brain areas of well-
educated subjects engaged in working memory tasks have less
small-world characteristics than those of less-educated volun-
teers. Iturria et al. (2008) estimated the anatomical connection
probabilities (ACP) between 90 cortical and subcortical brain
gray matter areas from diffusion-weighted Magnetic Resonance
Imaging (DW-MRI) techniques and concluded that all the
studied networks have small-world and broad-scale character-
istics. Van den Heuvel et al. (2008) used a voxel-wise approach
for a model-free examination of both inter-regional as well as
intra-regional in the human brain. Resting-state 3 Tesla fMRI
recordings of 28 healthy subjects were acquired and individual
connectivity graphs were formed out of all cortical and sub-
cortical voxels with connections reflecting inter-voxel function-
al connectivity. Graph characteristics from these connectivity
networks were computed. The clustering-coefficient of these
networks turned out to be much higher than the clustering-
coefficient of comparable random graphs, together with a short
average path length, indicating a small-world organization.
Furthermore, the connectivity distribution of the number of
inter-voxel connections followed a power-law scaling with an
exponent close to 2, suggesting a scale-free network topology.
Their findings suggested a combined small-world and scale-free
organization of the functionally connected human brain. The
results were interpreted as evidence for a highly efficient
209
organization of the functionally connected brain, in which
voxels are mostly connected with their direct neighbors forming
clustered sub-networks, which are held together by a small
number of highly connected hub-voxels that ensure a high level
of overall connectivity.
Appendix 3. The entropy as a measure of graph
connectivity
A graph G = ðN; A; WÞ is a mathematical description of a
network, consisting of a collection N of nodes n, A of arcs
a(ni, nj) connecting the nodes ni, nj and W of weights of
connections w(ni, nj)of a(ni, nj). The weight w(ni, nj) is a linear
function of the probability of the success of correctly transmit-
ting a signal from nitonj through the arc (channel) a(ni, nj).
Therefore w(ni, nj) → 1 when that probability tends to 1 and
conversely w(ni, nj) → 0 when the probability tends to 0. If signal
identity condition is relaxed by requiring the existence of
defined associations between the signals identified at ni and nj,
then the values of w(ni, nj) may be estimated by the correlation
between the observed signals at ni and nj, respectively. It may be
assumed that the correlation between the signals identified at
niandnjincreases as the probability of the success of correctly
transmitting a signal from nitonj increases. The average weight-
ing connection w ðni Þ of the node ni is calculated as:
c−1
∑ wðni ; nj Þ
j=1
w ðni Þ = ð1Þ
c−1
where c is the cardinality of G.
Appendix 1 provides a more detailed formal description of
the graph structure supporting small-world, scale-free and
broad-range networks. Its reading may be useful for the full
understanding of what follows.
Information is efficiently transmitted through small-world,
scale-free and broad range networks because the mean path
lengths of those network are small (ref). Following (Rocha
et al., 2004, 2005):
(a) the capacity of the channel supported by the arc a(ni, nj)
is given by the entropy h(ni, nj) defined as:
hðni ; nj Þ = −wðni ; nj Þlog2 wðni ; nj Þ−ð1−wðni ; nj ÞÞlog2 ð1−wðni ; nj ÞÞ;
ð2Þ
(b) the average amount of information handled by the node
ni if given by the entropy h ðni Þ calculated as:
h ðni Þ = −w ðni Þlog2 w ðni Þ−ð1−w ðni ÞÞlog2 ð1−w ðni ÞÞ; and ð3Þ
c) the net amount of information handled by the node ni if
given by the entropy h(ni) of the node ni, calculated as:
c−1
hðni Þ = ∑ ðh ðni Þ−hðni ; nj ÞÞ: ð4Þ
j=1
In this context, the actual value of h(ni) is:
(a) zero for any node of a totally connected graph because in
this case w(ni, nj) = 1 for all nj, and consequently h(ni, nj) = 0,
w ðni Þ = 1 and h ðni Þ = 0;
 Author's personal copy
210 BR A I N R ES E A RC H 1 3 5 1 ( 2 01 0 ) 1 9 8 –21 1
(b) zero for any node of a totally disconnected graph
because in this case w(ni, nj) = 0 for all nj, and conse-
quently h(ni, nj) = 0, w ðni Þ = 0 and h ðni Þ = 0;
(c) zero for any node of a random graph because in this case
w(ni, nj) = 0.5 for all nj, and consequently h(ni, nj) = 1,
w ðni Þ = 0:5 and h ðni Þ = 1.
In addition, in the case of the nodes ni ni highly connected
to c other nodes np, the actual value of h(ni):
c Hauk, O., Davis, M.H., Ford, M., Pulvermüller, F., Marslen-Wilson,
(d) tends to a value dependent on the ratio
n−1
w(ni, np) → 1 for all c highly connected nodes np and
w(ni, nq) → 0 for all the other remaining nodes nq = n − c,
such that n−1 c
≤w ðni Þ≤ 0:5nn−1
+ 0:5c
, 0 < h ðni Þ≤1, h(ni, np) → 0 and
0 ≤ h(ni, nq) ≤ 1.
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