International Journal of Psychophysiology 93 (2014) 322–331

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International Journal of Psychophysiology

journal homepage: www.elsevier.com/locate/ijpsycho

The effects of theta transcranial alternating current stimulation (tACS) on

fluid intelligence
Anja Pahor, Norbert Jaušovec ⁎
University of Maribor, Slovenia

a r t i c l e i n f o a b s t r a c t

Article history: The objective of the study was to explore the influence of transcranial alternating current stimulation (tACS) on
Received 1 March 2014 resting brain activity and on measures of fluid intelligence. Theta tACS was applied to the left parietal and left
Received in revised form 5 June 2014 frontal brain areas of healthy participants after which resting electroencephalogram (EEG) data was recorded.
Accepted 27 June 2014
Following sham/active stimulation, the participants solved two tests of fluid intelligence while their EEG was re-
Available online 3 July 2014
corded. The results showed that active theta tACS affected spectral power in theta and alpha frequency bands. In
Keywords:
addition, active theta tACS improved performance on tests of fluid intelligence. This influence was more pro-
Fluid intelligence nounced in the group of participants that received stimulation to the left parietal area than in the group of par-
tACS ticipants that received stimulation to the left frontal area. Left parietal tACS increased performance on the
Theta frequency difficult test items of both tests (RAPM and PF&C) whereas left frontal tACS increased performance only on the
EEG easy test items of one test (RAPM). The observed behavioral tACS influences were also accompanied by changes
ERD/ERS in neuroelectric activity. The behavioral and neuroelectric data tentatively support the P-FIT neurobiological
model of intelligence.
© 2014 Elsevier B.V. All rights reserved.

1. Introduction
Can we improve intelligence? The unanimous answer of researchers
is that, to this date, intelligence cannot be raised in a consistent and
long-lasting manner (Haier, 2014; Buschkuehl & Jaeggi, 2010; Jensen,
1998; Gottfredson, 1997). Recently, the debate on whether training
can increase performance on tests of intellectual ability has once more
gained popularity. The discussion was triggered by Jaeggi et al. (2008),
who showed that working memory (WM) training increased perfor-
mance on tests of fluid intelligence (Gf). Cognitive training, which typi-
cally targets working memory, reasoning, and executive functions, is the
most common form of intervention used to increase performance on
tests of intelligence (e.g., Owen et al., 2010; Klingberg, 2010; Morrison
& Chein, 2011; Chooi & Thompson, 2012; Colom et al., 2012;
Thompson et al., 2013). The reported effects are diverse, showing per-
formance increases (Jaeggi et al., 2008; von Bastian & Oberauer, 2013;
Stephenson & Halpern, 2013) or no significant effects (Owen et al.,
2010; Chooi & Thompson, 2012; Colom et al., 2012, 2013b; Thompson
et al., 2013). Some studies have further shown that these interventions
also change brain activation patterns (Jaušovec & Jaušovec, 2012).
Because our brain is the source of intelligent behavior (Kolb &
Whishaw, 2009), it could be speculated that interventions aimed
⁎ Corresponding author at: Univerza v Mariboru, Filozofska fakulteta, Koroška 160,
2000 Maribor, Slovenia. Fax: +386 2 258180.
E-mail address: norbert.jausovec@um.si (N. Jaušovec).
directly toward the structure and functional connectivity of the brain
would have a more pronounced influence on measures of fluid intelli-
gence. A direct approach might also have a methodological advantage
since it can rule out similarity effects between assessment and training
tasks and other environmental variables that are difficult to control or
exclude. Studies employing direct interventions can be divided into
three broad categories: neurofeedback training, pharmacological inter-
ventions, and brain stimulation. Several neurofeedback studies have
showed a modest positive influence on different tests of cognitive func-
tions such as episodic retrieving (Keizer et al., 2010), mental rotation
(Zoefel et al., 2011), music performance (Egner et al., 2004), creativity
and ballroom dancing (Gruzelier, 2009), and intelligence tests in a
group of individuals with an intellectual disability (Surmeli & Ertem,
2010). It has also been reported that nootropic or cognitive enhancing
substances modestly raise the performance of children on nonverbal
tests of intelligence (Schoenthaler et al., 2000), have a positive effect
on working memory and intelligence tests (Rae, et al., 2003), and in-
crease performance on Raven's Advanced Progressive Matrices
(Stough et al., 2011).
A third approach that can influence brain activity directly is the re-
cently rediscovered field of transcranial electrical stimulation (transcra-
nial direct current, and alternating current stimulation—tDCS and tACS).
It is assumed that tDCS modulates resting membrane potentials and, via
this mechanism, alters spontaneous cortical activity. Anodal tDCS en-
hances cortical activity and excitability, while cathodal stimulation has
the opposite effect (Bindman et al., 1964). On the other hand, tACS is a

http://dx.doi.org/10.1016/j.ijpsycho.2014.06.015
0167-8760/© 2014 Elsevier B.V. All rights reserved.
 A. Pahor, N. Jaušovec / International Journal of Psychophysiology 93 (2014) 322–331
newly developed stimulation technique that modulates cortical activity
by affecting neuronal membrane potentials via oscillatory electrical
stimulation in specific frequencies. This technique can be used to inter-
act with ongoing rhythmic cortical activities during cognitive processes
(Kuo & Nitsche, 2012). It has been suggested that tACS influences brain
oscillations via interference or entrainment of ongoing oscillations
(Thut & Miniussi, 2009), as demonstrated with rhythmic transcranial
magnetic stimulation (rTMS) in the alpha band (Thut et al., 2011). The
effectiveness of tACS in modulating oscillatory brain activity was also
demonstrated by Zaehle et al. (2010). Stimulation applied at partici-
pants' individual EEG alpha frequency (IAF) resulted in an enhancement
of the EEG alpha amplitude, indicating that this stimulation method can
interfere with ongoing brain oscillations in a frequency-specific manner.
In a recent study, Neuling et al. (2013) replicated and extended the find-
ings of Zaehle et al. (2010) by showing that the tACS-induced alpha am-
plitude enhancement remained present for at least 30 min after
stimulation offset.
To our knowledge, just one study has been conducted that investi-
gates the influence of transcranial electrical stimulation on tests of
fluid intelligence. Santarnecchi et al. (2013) showed that gamma-band
tACS delivered over the left middle frontal gyrus resulted in a shorten-
ing of the time required to find the correct solution in a visuospatial ab-
stract reasoning test similar to Raven's matrices (1990). However,
reaction time may not serve as a valid indicator of fluid intelligence, es-
pecially since the standard application of Raven's matrices has no time
limitation. On the other hand, several studies have reported positive in-
fluences of tDCS and tACS on tests of cognitive processes related to fluid
intelligence, such as working memory performance (Mulquiney et al.,
2011; Boggio et al., 2008; Fregni et al., 2006; Polanía et al., 2012;
Tseng et al., 2012), the ability to discriminate numerosity (Cappelletti
et al., 2013), creativity, and problem solving (Cerruti & Schlaug, 2009;
Chi & Snyder, 2011; Dockery et al., 2009; for a review see Kuo &
Nitsche, 2012).
The aim of the present study was to explore the influence of dif-
ferent tACS protocols on performance on tests of fluid intelligence
and to extend the current understanding of the neuronal underpin-
nings of intelligence. Fluid intelligence is defined as the capacity to
solve novel, complex problems, using operations such as inductive
and deductive reasoning, concept formation, and classification. It
has also been suggested that Gf represents the influences of biologi-
cal factors and incidental learning on intellectual development
(Cattell, 1971; Horn & Cattell, 1966). Even though the influence of
tDCS on higher cognitive functions has been explored to a greater ex-
tent than the influence of tACS, we decided to use tACS because of the
possibility to investigate the relationship between brain oscillations
and performance on tests of fluid intelligence. Synchronized cortical
oscillations in different frequency bands have been proposed to be
an important mechanism of high-level cognition giving rise to intel-
ligence, for example, theta band activity has been linked to working
memory while narrow alpha band frequencies have been related to
attention (Klimesch, 2012).
With respect to the waveform and the specific oscillatory frequency
used in the present study, two aspects were considered: (1) the
established relation between the constructs of WM and intelligence re-
ported in correlational (Buehner et al., 2005; Colom et al., 2008;
Oberauer et al., 2008; Martínez et al., 2011) and experimental ap-
proaches (for a review see Morrison and Chein, 2011; Rabipour & Raz,
2012); and (2) the relation between working memory processes and
neuronal rhythms in the theta band. It has been shown that theta syn-
chronizes during working memory processes and serves as a gating
mechanism, providing optimal neural conditions for specific processing
(Sauseng et al., 2010). Therefore, theta oscillation was applied in the
tACS protocol. In particular, our goal was to examine whether theta
tACS can affect power in theta and alpha frequency bands during a sub-
sequent period of rest and during performance on tests of fluid
intelligence.
323
The second tACS protocol variable that was determined prior to
conducting the study was location of stimulation (placement of active
electrodes). Based on research findings concerning the structure and
neuronal origin of intelligence we decided to target left frontal and left
parietal brain areas. According to a model proposed by Duncan et al.
(2000), psychometric g is implemented within the prefrontal cortex,
whereas a more recent model claims that intelligence depends on dis-
tributed system of functionally specialized fronto-parietal cortical re-
gions (Jung & Haier, 2007). In Duncan's model (Duncan, 2001, 2003;
Duncan & Owen, 2000) fluid intelligence mainly derives from a specific
frontal brain network (mid-dorsolateral, mid-ventrolateral and dorsal
anterior cingulate cortex), which is important for the control of diverse
cognitive functions (e.g., executive control, strategy formation, monitor-
ing). Some neuroscience data support this framework, demonstrating
the involvement of the prefrontal cortex in performance on tests of in-
telligence (Roca et al., 2010; Bishop et al., 2008; Gray et al., 2003;
Prabhakaran et al., 1997) and executive function (Duncan, 2006;
Duncan & Owen, 2000). A recent morphometric study further indicated
that structural features of gray matter in the frontal lobes show an over-
lapping cluster with measures of crystallized and fluid intelligence
(Colom et al., 2013a). On the other hand, Jung and Haier (2007)
reviewed 37 neuroimaging studies on the structural correlates of intel-
ligence and synthesized their findings into a so-called ‘parieto-frontal
integration’ (P-FIT) model of intelligence. This neurobiological model
of intelligence emphasizes the importance of frontal and parietal re-
gions and the white matter association tracts that bind these areas
into a synchronized system (Jung & Haier, 2007). Indeed, a number of
studies have demonstrated the importance of parietal regions in perfor-
mance on tests of general intelligence with the use of neuroimaging
techniques (Lee et al., 2006; Colom et al., 2006; Haier et al., 2004), elec-
troencephalography (Jaušovec & Jaušovec, 2004; Gevins & Smith, 2000),
and lesion mapping (Barbey et al., 2012; Gläscher et al., 2009).
Based on previous research on the effects of IAF tACS on alpha power
(Zaehle et al., 2010; Neuling et al., 2013) we hypothesized that active
theta tACS would increase theta power, but would have no effect on fre-
quencies in the alpha range. Furthermore, we predicted that theta tACS
would improve performance on tests of fluid intelligence. In contrast, no
improvement in performance would be seen following sham tACS. In-
creased performance on IQ tests induced by theta tACS would suggest
brain activation patterns similar to those observed in highly intelligent
individuals. In two EEG studies (Gevins & Smith, 2000; Jaušovec &
Jaušovec, 2004) as well as in one fMRI study (Rypma et al., 2006) indi-
viduals scoring high on IQ tests displayed less frontal but more parietal
cortical activation. Thus, it was predicted that active tACS positioned
over parietal brain areas would have a more prominent effect on perfor-
mance on tests of fluid intelligence than active tACS positioned over
frontal brain areas.
2. Method
2.1. Subjects
Individuals participating in the experiment were recruited from a
large scale resting (eyes-closed) EEG study. They were selected because
their eyes-closed individual alpha peak frequency (IAF), which is need-
ed to determine theta frequency for tACS, and their IQ scores (WAIS-R,
Wechsler, 1981), were available. The sample included 28 right-handed
individuals (20 females; average age = 20 years and 8 months; SD =
4.35 months) that were assigned to two groups and equalized with re-
spect to IQ: Parietal group (N = 14; MIQ = 105.38, SD = 9.25) and
Frontal group (N = 14; MIQ = 105.45; SD = 8.93). The participants
had a similar educational background, took no medication, and reported
no health problems. Due to artifacts in the resting (eyes-closed) EEG
data, 4 individuals were excluded from the resting EEG data analysis
(N = 24; 16 females; average age = 20 years and 7 months; SD =
5.25 months).
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2.2. Tasks and procedure
The individuals participated in two sessions—sham and active tACS,
which were counterbalanced. The sham and active settings were sepa-
rated by 28 days in order to ensure that the females were tested on
the same day of their menstrual cycle. It has been shown that the rela-
tive release of sexual hormones during different phases of the menstrual
cycle affects cognitive responses of females (Amin et al., 2006). Specifi-
cally, high levels of estrogen and progesterone have been associated
with positive affect and higher prefrontal cortex activity. Fig. 1A shows
the timeline of the experiment. During both sessions, the participants
were exposed to 15 min of tACS (in the sham setting only for 1 min—
see description below) after which they rested with their eyes closed
for 5 min while their EEG was recorded. Subsequently they solved two
tests of fluid intelligence (RAPM and PF&C) during which EEG data
was also collected. At the end of the experiment the participants an-
swered a questionnaire about their sensations during the stimulation
period.
The RAPM test was based on a modified version of Raven's progres-
sive matrices (Raven, 1990), a widely used and well established test of
fluid intelligence (Sternberg et al., 1996). The correlation between this
modified version of RAPM and WAIS-R was r = .56 (p b .05, N = 97)
(Jaušovec & Jaušovec, 2012). Similar correlations of the order of 0.40
to 0.75 were also reported for the standard version of RAPM (Court &
Raven, 1995). For this reason it can be assumed that the modified appli-
cation of the RAPM did not significantly alter its metric characteristics.
The task was presented on a computer screen, which was positioned ap-
proximately 80–100 cm away from the participant. In a given trial, a
Fig. 1. (A) Sequence of the experiment. (B) Procedure of the PF&C test, which also applies to the RAPM test. (C) Experimental setup: tACS target electrodes positioned over left frontal (F3)
or left parietal sites (P3) and a return electrode positioned above the right eyebrow (Fp2) of the 10/20 system.
figural matrix with the lower right entry missing appeared on the
screen and the participant was asked to determine which of 4 different
options fits into the blank space. The test items were exposed for 6 s
(easy) or 10 s (difficult) at an interstimulus interval of 10 or 14 s respec-
tively. Following the presentation of the item and a 2 s interval, a cross
was presented during which the participant was instructed to press a
button on a response pad (1–4) to indicate his or her answer
(Jaušovec & Jaušovec, 2012). We used 50 test items—25 easy (Advanced
Progressive Matrices Set I: 12 items and the B Set of the Colored Progres-
sive Matrices) and 25 difficult (Advanced Progressive Matrices Set II:
items 12 to 36). The 50 test items were divided into 2 parallel forms
consisting of 25 items each that were used interchangeably for sham
and active conditions. The correlation between the two equivalent test
forms was r = 0.73 (p b .01, N = 120).
The second task used was based on the Paper Folding and Cutting
(PF&C) subtest of the Stanford–Binet IQ test (Rideout & Laubach,
1996). According to the authors, the subtest measures visual–spatial
reasoning. In a given trial, the left part of a computer screen showed
how a piece of paper was folded and cut while the right side of the
screen showed four unfolded papers, one of which corresponded to
the folded paper on the left. The participants were asked to judge
which of the four figures on the right corresponded to the figure on
the left (Jaušovec & Jaušovec, 2012). On sham/active settings, two paral-
lel forms consisting of 20 test items were used. The correlation between
the two equivalent test forms was r = 0.71 (p b .01, N = 120). The pre-
sentation was the same as that of the RAPM except that the 20 items
were exposed for 7 s. Fig. 1B shows the timeline of the PF&C task,
which roughly corresponds to the timeline of the RAPM task. The
 A. Pahor, N. Jaušovec / International Journal of Psychophysiology 93 (2014) 322–331
correlation between performance on RAPM and PF&C tests in this study
compared to performance on the same tests in a previous study
(Jaušovec & Jaušovec, 2012) was r = 0.43 (p b .01; N = 122).
2.3. Electrical stimulation
Transcranial alternating current stimulation (tACS) was applied via
two sponge electrodes attached to the head underneath an EEG record-
ing cap (Quik-Cap Compumedics Neuromedical supplies, Charlotte, NC,
USA). Fig. 1C shows the placement of the electrodes: the target electrode
was placed either over the left parietal location (P3) or the left frontal lo-
cation (F3), while the return electrode was placed above the right
orbitofrontal cortex (Fp2)—above the right eyebrow. The electrodes
were placed in a saline soaked sponge. To increase the focus of stimula-
tion, the size of the target electrode was 7 × 5 cm whereas the size of the
reference electrode was 10 × 7 cm (Moliadze et al., 2010). The waveform
of the stimulation was sinusoidal without DC offset with a 0° relative
phase. The impedance was kept below 10 kΩ. We applied oscillating
theta currents based on the IAF of each participant (theta = IAF-5 Hz;
M = 5.07; SD = 1.25) using a battery-operated stimulator system
(DC-stimulator plus, Neuroconn, Ilmenau, Germany). In the active con-
dition, tACS was applied for 15 min. The current was ramped up and
down over the first and last 15 s of stimulation. In the sham condition,
the procedure was the same except for the duration of stimulation,
which was applied for 1 min and then turned off automatically. Because
most participants feel an itching sensation only initially during tACS,
this procedure prevents awareness of the stimulation conditions
(Nitsche et al., 2008).
The magnitude of the stimulating current was based on individually
determined thresholds for skin sensations induced by tACS. To determine
these thresholds, we applied tACS stimulation at the individual theta fre-
quency for 1 min (ramping up and down for 15 ms) at a time and in-
creased the amplitude stepwise by 250 μA starting with 1000 μA until a
maximum of 2250 μA was reached. Participants were asked to keep
their eyes closed and to indicate the presence of a sensation. For the re-
maining experiment, stimulation intensity was kept at 250 μA below
the threshold for skin sensations (Modus = 1750 μA peak-to-peak;
Range = 1000 μA to 2250 μA). The tACS parameter characteristics were
in the range suggested by Zaghi et al. (2010) for the (1) stimulation du-
ration 5 min–30 min, (2) stimulation intensity 500 μA–2000 μA, and
(3) scalp current density 24 μA/cm2–29 μA/cm2.
2.4. EEG recording
EEG was recorded using a Quick-Cap with sintered (Silver/Silver
Chloride; 8 mm diameter) electrodes. Using the 10–20 Electrode Place-
ment System of the International Federation, EEG activity was moni-
tored over 19 scalp locations (Fp1, Fp2, F3, F4, F7, F8, T3, T4, T5, T6,
C3, C4, P3, P4, O1, O2, Fz, Cz, and Pz). All leads were referenced to linked
mastoids (A1 and A2), and a ground electrode was applied to the fore-
head. Additionally, vertical eye movements were recorded with elec-
trodes placed above and below the left eye. Electrode impedance was
maintained below 5 kΩ. The digital EEG data acquisition and analysis
system (SynAmps RT) had a bandpass of 0.15–100.0 Hz. At cutoff fre-
quencies, the voltage gain was approximately −6 dB. The 19 EEG traces
were digitized online at 1000 Hz with a gain of 10 × (accuracy of
29.80 nV/LSB in a 24 bit A to D conversion) and stored on a hard disk.
2.5. Resting EEG data analysis
In order to examine whether sham and active stimulation led to
different baseline activity, resting EEG data was analyzed in terms
of spectral power. Comprised were epochs of 4096 data points. A
Fast Fourier Transformation (FFT) was performed on the epoched
EEG data in order to derive estimates of absolute spectral power
(uV) in different frequency bands. The frequency bands were
325
individually determined in 2.0 Hz steps based on individual alpha
peak frequency (MIAF = 10.07; SDIAF = 1.25) for four bands: upper
alpha = [IAF to IAF + 2.0 Hz]; lower-2 alpha = [IAF to IAF − 2.0 Hz];
lower-1 alpha = [IAF − 2.0 Hz to IAF − 4.0 Hz]; and theta =[IAF
− 4.0 Hz to IAF − 6.0 Hz] (Klimesch, 1999).
2.6. Task-related EEG data analysis
For the EEG data recorded during task performance, epochs were ex-
tracted, ranging from 2000 ms before stimulus presentation to 10,000 ms
after its presentation. Prior to the analysis, a correction for ocular artifacts
was performed and the epochs were automatically screened for artifacts.
All epochs showing amplitudes above ±80 μV were excluded. Overall,
2.1% of epochs were excluded from further analysis. The induced
event-related desynchronization/synchronization (ERD/ERS) was deter-
mined using the method of complex demodulation with a simultaneous
signal envelope computation (Pfurtscheller, 1999). For this method, raw
data for each channel were multiplied, point-by-point, by a pure cosine
based on the selected center frequency and by a pure sine with the
same center frequency. Both time series (multiplied by a pure sine and
cosine) were then low pass filtered by the half-bandwidth (1 Hz).
The quantification of induced ERD was done using the inter-trial var-
iance method (induced, non-phase-locked activity). The formulas used
were as follows (Pfurtscheller, 1999):
N n o2
1 X
IV ð jÞ ¼ y f ði; jÞ −y f ð jÞ : ð1Þ
N−1 i¼1
In Eq. (1) N is the total number of trials, yf(i,j) is the j-th sample of i-th
trial data, and y f ð jÞ is the mean of the j-th sample over all trials. ERD (IV)
data were used to calculate the ERD/ERS values that were defined as the
percentage change of the power at each sample point (Aj) relative to the
average power during the resting 1000 ms reference interval
(R) preceding the stimulus onset (−1500 ms to −500 ms):
R−Aj
ERD=ERSð jÞ % ¼ : ð2Þ
R
A positive ERD indicates a power decrease, and a negative ERS indi-
cates a power increase (Pfurtscheller, 1999). Due to statistical con-
straints, we did not conduct separate ERD/ERS analyses for easy and
difficult RAMP items. The ERD/ERS values were determined for six
1000 ms time windows (from stimulus onset till 6000 ms) and were
collapsed for different electrode locations, distinguishing the hemi-
spheres as well as frontal, central, and parietal brain areas. The electrode
positions were aggregated as follows: frontal left (Fp1, F3, F7, Fz), fron-
tal right (Fp2, F4, F8, Fz), central left (T3, C3, Cz), central right (T4, C4,
Cz), parietal left (T5, P3, O1, Pz), and parietal right (T6, P4, O2, Pz).
3. Results
3.1. Resting EEG
To explore the influences of tACS on power measures, a GLM for re-
peated measures was conducted separately for the Parietal and the Fron-
tal group. The following variables were defined: tACS (sham/active),
hemisphere (left/right), and location (frontal, central, parieto-occipital).
In the Frontal group, the only significant influence of theta tACS on
power measures was observed in the lower-2 alpha band. The GLM
showed a significant interaction effect between tACS and location
(F(2,22) = 3.80; p = 0.04; partial eta2 = 0.26). Fig. 2 shows that the de-
crease in lower-2 alpha power was most prominent in frontal brain areas.
The GLM for the Parietal group showed significant differences be-
tween sham and active settings in theta, lower-1, and lower-2 alpha fre-
quency bands, all showing a significant interaction between tACS and
location (theta: F(2,22) = 5.90; p = 0.009; partial eta2 = 0.35; lower-
326 A. Pahor, N. Jaušovec / International Journal of Psychophysiology 93 (2014) 322–331

Fig. 2. (A) Topographic power maps of sham and active resting eyes-closed neuroelectric activity in the theta band and (B) alpha band (collapsed for the lower-1 and lower-2 alpha bands)
after theta tACS—target electrode positioned over the left parietal site (P3).

1 alpha: F(2,22) = 7.79; p = 0.003; partial eta2 = .42; lower-2 alpha: On the other hand, in the two alpha bands, participants in the active set-
F(2,22) = 11.10; p = 0.001; partial eta2 = 0.50). As can be seen in ting showed decreased alpha power mainly in the posterior brain areas
Fig. 3A, in the active setting participants showed increased theta (Fig. 3B). Note that in Fig. 3B the power spectra for the lower-1 and
power mainly in left frontal brain areas compared to the sham setting. lower-2 alpha frequency bands are collapsed.

Fig. 3. Topographic power map of sham and active resting eyes-closed neuroelectric activity in the lower-2 alpha band after theta tACS—target electrode positioned over left frontal site
(F3).
 A. Pahor, N. Jaušovec / International Journal of Psychophysiology 93 (2014) 322–331 327

3.2. Behavioral data
The questionnaire data were analyzed with a Wilcoxon nonpara-
metric test for two related samples (sham/active). The test showed
no significant differences between participants' sensations during
sham and active tACS (Z(11) = .06; p = 0.98). Performance data
were analyzed with a general linear model (GLM) for repeated
measures—tACS (sham/active) × task (RAPM/PF&C) × group (Parie-
tal/Frontal). The GLM showed a significant main effect of tACS
(F(1,26) = 27.19; p b .001; η2 = .51), and a significant interaction ef-
fect between tACS and group (F(1,26) = 6.19; p = .02; η2 = .19). Ac-
tive tACS improved performance on tests of fluid intelligence. This
influence was more pronounced for the Parietal group. Subsequent
paired-sample t tests between sham and active settings, which were
FDR-corrected (Benjamini & Hochberg, 1995), were significant both
for RAPM and PF&C tests. These differences were more pronounced
for the difficult test items than for the easy items. In the active setting,
participants scored notably higher on both tests, mainly by correctly
solving more difficult test items than in the sham setting.
Participants in the Frontal group also showed a significant increase
in performance, however, subsequent paired sample t-tests between
the sham and active settings for the RAPM and PF&C total test scores
were not significant. As can be seen in Table 1, the only significant differ-
ence between sham/active settings observed for this group was for the
easy RAPM test items. In the active setting the participants correctly
solved more easy RAPM test items than in the sham setting.
3.3. Task-related EEG
In order to examine whether the site of stimulation influenced oscil-
latory power during performance on tests of fluid intelligence, a GLM for
repeated measures with the following factors was conducted: tACS
(sham/active) × task (RAPM/PF&C) × time (six 1000 ms segments) ×
hemisphere (left/right) × location (frontal, central, occipito-parietal) ×
group (Parietal/Frontal). A significant effect of the factor group was ob-
served in the theta, lower-1 and lower-2 alpha frequency bands. In the
theta frequency band, the GLM analysis showed a significant interaction
effect among thevariables tACS, task, and group (F(1,26) = 8.00; p =
.009; η2 = .24); and among the variables tACS, location, and group
(F(2,52) = 4.51; p = .02; η2 = .15). In the lower-1 alpha frequency
band, a significant interaction effect among the variables tACS, task,
hemisphere and group (F(1,26) = 4.30; p = .04; η2 = .14) was observed.
In the lower-2 alpha frequency band a significant main effect of tACS
(F(1,26) = 6.87; p = .01; η2 = .21); and an interaction effect among
Table 1
Descriptive statistics (means and standard deviations) for RAPM and PF&C tests (distin-
guished between total scores and scores on easy/difficult items) in sham and active tACS
settings for the Parietal and Frontal groups.
the variables tACS, location and group (F(2,52) = 4.39; p = .02; η2 =
.14) were observed.
To gain a deeper insight into the influences of tACS on brain activa-
tion patterns during problem solving, a GLM was conducted separately
for each group: tACS (sham/active) × task (RAPM/PF&C) × time (six
1000 ms segments) × hemisphere (left/right) × location (frontal, cen-
tral, occipito-parietal).
In the Frontal group, active compared to sham tACS significantly af-
fected power in two bands: theta and lower-1 alpha. Namely, in the
theta band, significant interaction effects between the variables tACS
and task (F(1,13) = 4.36; p = .05; η2 = .25); tACS and location
(F(2,26) = 5.34; p = .01; η2 = .29); and among the variables tACS,
task, and time (F(5,55) = 2.88; p = .04; η2 = .21) were observed. Dif-
ferences in theta ERD/ERS between active and sham settings were more
pronounced for the RAPM test. Fig. 4 shows that in the active setting,
participants displayed a time related (3000–6000 ms) decrease in
theta power mainly in frontal brain areas while solving the RAPM task.
In the lower-1 alpha band, there was a significant interaction effect
among tACS, task, and hemisphere (F(1,13) = 8.77; p = .01; η2 = .40).
As can be seen in Fig. 5, during the RAPM test, participants in the active
setting showed a decrease in lower-1 alpha power over the left hemi-
sphere compared to the sham setting.
In the Parietal group, significant differences in power as a factor of
tACS were observed only in one band: lower-2 alpha. Specifically, a sig-
nificant main effect of tACS (F(1,13) = 14.45; p = .002; η2 = .53),
and an interaction effect between the variables tACS and location
(F(1,13) = 4.19; p = .03; η2 = .24) were observed. As can be seen in
Fig. 6, participants in the active setting showed higher lower-2 alpha
ERD than in the sham setting, which was more pronounced in frontal
brain areas.
4. Discussion
The results showed that active tACS in the theta frequency signifi-
cantly influenced resting brain oscillatory activity. Theta tACS applied
to left frontal and parietal areas significantly decreased lower-1 and
lower-2 alpha power in brain areas located close to or beneath the site
of stimulation. Furthermore, theta tACS delivered to the left parietal
area significantly increased theta power in frontal brain areas. The effect
of active tACS compared to sham stimulation was also observed on per-
formance on tests of fluid intelligence. Active theta tACS improved per-
formance on Raven's progressive matrices and on the Paper Fold & Cut
test. The improvement was more pronounced in the group of partici-
pants that received stimulation to the left parietal area than in the
group that received stimulation to the left frontal area. Spectral power
calculated during performance on these tests showed that the parietal
stimulation had a general influence on ERD/ERS patterns, whereas fron-
tal stimulation showed a task dependent influence that was related to
improved performance on the RAPM test.

Sham setting Active setting df(13) 4.1. tACS effects on resting EEG
M SD M SD
The observed effects of theta tACS on resting neuronal electric activ-
Parietal
RAPM—total 13.50 3.35 15.79 2.01 t = 2.88; p = .01⁎; d = .77 ity demonstrate that tACS can be used to specifically modulate oscillato-
RAPM—easy 8.86 1.70 9.64 .63 t = 1.76; p = .10; d = .47 ry activity in particular brain areas—most likely by interfering with the
RAPM—difficult 4.64 2.27 6.14 1.92 t = 3.07; p = .009⁎; d = .82 ongoing rhythmic activity of local pacemaker networks. Interestingly,
PF&C—total 10.29 2.13 12.64 3.43 t = 3.16; p = .008⁎; d = .77
the after-effects of theta tACS were more prominent when stimulation
PF&C—easy 4.93 1.27 5.57 1.79 t = 1.04; p = .41; d = .28
PF&C—difficult 5.36 1.87 7.07 1.92 t = 3.97; p = .002⁎; d = 1.06
was applied to the left parietal site compared to the left frontal site. Pa-
rietal stimulation led to an increase in theta power in left frontal areas
Frontal
and a decrease in lower-1 alpha and lower-2 alpha power in posterior
RAPM—total 14.00 2.72 14.86 3.13 t = 1.06; p = .18; d = .28
RAPM—easy 8.71 1.44 9.72 1.33 t = 2.39; p = .03⁎; d = .64
areas. The functional importance of stimulating in theta frequency
RAPM—difficult 5.29 1.68 5.14 2.63 t = .22; p = .83; d = .06 should be considered in the light of the findings that (1) local theta ac-
PF&C—total 11.71 2.27 12.50 3.16 t = 1.01; p = .31; d = .27 tivity increases during working memory processes, and (2) interregional
PF&C—easy 5.86 1.17 5.36 1.15 t = 1.29; p = .22; d = .34 synchronization of theta oscillations connects different neural struc-
PF&C—difficult 5.85 1.56 7.14 2.88 t = 1.66; p = .12; d = .44
tures that are involved in working memory processes (Sauseng et al.,
⁎ FDR adjusted significance. 2010). The finding that theta tACS applied to the left parietal area
328 A. Pahor, N. Jaušovec / International Journal of Psychophysiology 93 (2014) 322–331

Fig. 4. Topographic power map of neuroelectric activity in the theta frequency band recorded during the RAPM test after sham and active theta tACS—target electrode positioned over the
left frontal site (F3).

increased theta power in frontal areas supports the idea that theta oscil-
lations reflect a general integrative mechanism in the brain (Sauseng
et al., 2010). On the other hand, tACS applied to the left frontal area re-
sulted in a decrease in lower-2 alpha power in frontal areas, but did not
affect theta power or any other alpha bands. Stimulating the left parietal
region might have had a more prominent effect on brain activity due
to the fact that it has been identified as one of the brain regions
(termed hubs) that plays a central role in the organization of neural net-
works (Van den Heuvel and Sporns, 2013; Langer et al., 2012; Achard
et al., 2006).

Fig. 5. Topographic power map of neuroelectric activity in the lower-1 alpha frequency band recorded during the RAPM test after sham and active theta tACS—target electrode positioned
over the left frontal site (F3). The power maps are averaged over 6000 ms.
 A. Pahor, N. Jaušovec / International Journal of Psychophysiology 93 (2014) 322–331
Fig. 6. Topographic power map of neuroelectric activity in the lower-2 alpha frequency band recorded during the RAPM and PF&C tests after sham and active theta tACS—target electrode
positioned over the left parietal site (P3). The power maps are averaged over 6000 ms.
4.2. Behavioral results
Stimulation applied to the left parietal brain area had a more pro-
nounced effect on performance on tests of fluid intelligence than stimu-
lation applied to the left frontal area. This finding is in line with research
demonstrating the relationship between performance on tests of intel-
ligence and brain activity in parietal areas (Lee et al., 2006; Haier et al.,
2003; Wharton et al., 2000) and lends further support to the assump-
tion that working memory storage capacity largely accounts for the re-
lationship between the psychological constructs of working memory
and intelligence (Colom et al., 2008; Buehner et al., 2005). The impor-
tance of the left parietal lobe for working memory storage capacity
has been demonstrated by several neuroimaging (Cowan et al., 2011;
Cowan, 2011; Majerus et al., 2010; Xu & Chun, 2006; Todd & Marois,
2004), rTMS (Postle et al., 2006), and tDCS (Tseng et al., 2012) studies.
In a recent study by Jaušovec and Jaušovec (2014), theta tACS applied
to the left parietal area significantly increased participants' working
memory capacity. Furthermore, the ERP analysis indicated a decrease
in P300 latency suggesting an improvement in the participants' ability
to rapidly allocate attention resources needed to solve the task. A similar
finding was also reported by Tseng et al. (2012), who showed that tDCS
over the right parietal cortex mainly improved the ability to suppress ir-
relevant distractors in poor performers.
On the other hand, stimulating the left frontal area had a minor effect
on test performance. On the RAPM test, participants scored higher on
easy test items in the active stimulation session compared to the sham
session. There were no differences in performance on the PF&C test be-
tween the two sessions. This finding is in line with research showing
that low-ability subjects rely more exclusively on frontal regions during
performance on tests of intelligence, whereas high-ability individuals
make relatively greater use of parietal regions (Lee et al, 2006;
Jaušovec & Jaušovec, 2004; Gevins & Smith, 2000). Furthermore,
Langer et al. (2012) demonstrated that the more intelligent a person
is, the more his or her brain networks resemble a small world network.
In particular, the parietal cortex was identified as a main hub within
such small world networks.
The analysis of the influence of tACS on easy and difficult test items
revealed that left parietal tACS had a more pronounced influence on dif-
ficult test items (RAPM and PF&C), whereas left frontal tACS had a more
pronounced influence on easy test items (RAPM). It can be assumed that
329
stimulation in parietal and frontal areas differently influenced cognitive
processes involved in fluid intelligence. One possible explanation may
be that left parietal tACS mainly increased WM storage capacity thereby
improving performance on difficult test items, whereas left frontal tACS
mainly influenced the attentional components required for successful
performance on easy test items. Support for the former interpretation
comes from the relation between working memory processes and neu-
ronal rhythms in frontal and parietal brain areas (Sauseng et al., 2010)
and from studies employing tACS (Jaušovec & Jaušovec, 2014;
Jaušovec et al., 2014). Support for the latter claim comes from a recent
study in which active tDCS applied to the prefrontal cortex resulted in
an improvement in target detection performance (Nelson et al., 2014).
According to the authors, active tDCS enhanced sustained attention,
which improved task performance. However, given the fact that a grow-
ing number of studies consider working memory and attention as over-
lapping constructs (Gazzaley & Nobre, 2012; Chun, 2011; Postle, 2006;
Awh et al., 2006), specific conclusions about the effects of tACS on
task-related mental processes cannot be made.
4.3. Task-related EEG
The observed effects of active tACS on behavioral variables were also
accompanied by changes in neuroelectric activity during performance
on tests of fluid intelligence. Active stimulation applied to the left pari-
etal area was associated with a global decrease in lower-2 alpha power
in the whole brain, which was the strongest in left frontal areas. Re-
search suggests that a reduction of power in lower alpha reflects general
task demands, particularly attentional processes (Klimesch et al., 1999;
Doppelmayr et al., 2002). In contrast, frontal stimulation was associated
with a task-dependent decrease in theta power in left frontal areas
along with improved performance on easy RAPM test items. During
the RAPM test, participants in the active setting also showed a decrease
in lower-1 alpha power over the left hemisphere. Active theta tACS ap-
plied to the left frontal area either increased the participants' focus of at-
tention or improved binding in working memory, which resulted in
improved performance on the easy RAPM test items.
There are two prominent neurobiological models that explain the
relation between intelligence and brain activity: the parieto-frontal in-
tegration (P-FIT) model (Jung and Haier, 2007) and Duncan's adaptive
coding model (Duncan, 2001, 2003). On the whole, the results of the
330 A. Pahor, N. Jaušovec / International Journal of Psychophysiology 93 (2014) 322–331
present study lend support to the P-FIT model proposed by Jung and
Haier (2007), suggesting that discrete brain regions play a role in gener-
al intelligence, with an emphasis on the parietal cortex and do not lend
support to Duncan's (2003) assumption that psychometric g is entirely
executed within processing networks of the prefrontal cortex.
The main limitation of the current study is that we did not record
pre-stimulation resting EEG data. Comparing pre- and post-stimulation
EEG data would have given further insight into the effects of theta tACS
on brain activity. Another shortcoming of the study is the predominate-
ly female structure of the sample. Given the immense body of evidence
with regard to sex related differences in general, as well as differences in
specific abilities (e.g., Johnson & Bouchard, 2007; Nyborg, 2005), the
structure of the sample might have influenced the results. Furthermore,
tACS was delivered only in the theta frequency band, therefore it cannot
be assumed that the obtained results are specific for theta stimulation.
Nevertheless, the results of the study demonstrate for the first time
that theta tACS increases resting theta power and, contrary to existing
research (Zaehle et al., 2010; Merlet et al., 2013), also affects power in
frequencies other than the stimulating frequency, namely in the alpha
band. Moreover, the results showed that theta tACS applied to the left
parietal brain area moderately increases performance on tests of fluid
intelligence. The observed behavioral and neuroelectric data tentatively
support the neurobiological P-FIT model of intelligence. Future research
could examine whether theta tACS combined with working memory
training further improves performance on tests of fluid intelligence.
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