Neurobiology of Learning and Memory 157 (2019) 114–120

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Neurobiology of Learning and Memory

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

Transcranial direct current stimulation (tDCS) over parietal cortex improves T

associative memory
Jovana Bjekića, , Marija V. Čolića, Marko Živanovićb, Sladjan D. Milanovića, Saša R. Filipovića
⁎

a
Department of Neuroscience, Institute for Medical Research, University of Belgrade, Serbia
b
Department of Psychology, Faculty of Philosophy, University of Belgrade, Serbia

ARTICLE INFO ABSTRACT

Keywords: Associative memory plays a key role in everyday functioning, but it declines with normal ageing as well as due to
Transcranial direct-current stimulation (tDCS) various pathological states and conditions, thus impairing quality of life. Associative memory enhancement via
Posterior parietal cortex (PPC) neurostimulation over frontal areas resulted in limited success, while posterior stimulation sites seemed to be
Cognitive enhancement more promising. We hypothesized that anodal transcranial direct current stimulation (tDCS) of parietal areas
Associative memory
would lead to higher performance in associative memory due to high connectivity between posterior parietal
Face-word memory
Object-location memory
cortex (PPC) and hippocampus. Forty-two healthy adults participated in two sham-controlled cross-over ex-
periments. Anodal electrode (20 min, 1.5 mA) was placed over P3 in Experiment 1 and over P4 in Experiment 2.
During tDCS participants played a simple computer game. After each stimulation session, participants completed
parallel forms of an associative memory task (Experiment 1: face-word memory; Experiment 2: object-location
memory) and a control task (verbal fluency). In both experiments, associative memory was improved after
anodal stimulation compared to sham stimulation, while no differences were observed in the control task.
Additionally, memory performance was higher in the second than in the first trial, but the increase in perfor-
mance between the two trials did not differ between stimulation conditions. It can be concluded that a single-
session anodal tDCS over posterior parietal cortex can improve associative memory performance. The specificity,
robustness, and reproducibility of the effect suggest that PPC is a promising target for brain stimulation aiming
to enhance memory functions.

1. Introduction
Associative or relational memory refers to the ability to learn and
remember the relationship between different and previously unrelated
pieces of information in same (e.g. visual: object-location associations)
or different modalities (e.g. visual and verbal: face-name associations)
(Konkel & Cohen, 2009). This type of memory plays a crucial role in
everyday functioning, but it is one of the first to be affected by different
neurological disorders (Giovanello, Verfaellie, & Keane, 2003), as well
as by normal (Chalfonte & Johnson, 1996) and pathological ageing
(Bastin et al., 2014). Therefore, the ability to improve associative
memory performance presents one of the most challenging tasks in
cognitive enhancement and neurorehabilitation.
Over the past decade, transcranial direct current stimulation (tDCS)
has become one of the most widely used techniques for non-invasive
brain stimulation. The tDCS modulates cortical excitability by deli-
vering weak electrical current between electrodes positioned over the
scalp. It is generally assumed that anodal stimulation facilitates
processes associated with the cortical region under the target electrode,
whereas cathodal stimulation has the opposite effect (Medeiros et al.,
2012). Facilitatory effects of anodal tDCS have been demonstrated
across multiple motor and cognitive domains in both healthy and
clinical populations (Kuo & Nitsche, 2012). The after-effects of tDCS are
considered to be NMDA-receptor dependent, and similar to the long-
term potentiation process that underlies learning and memory
(Medeiros et al., 2012). Still, the current understanding of the exact
physiological consequences of tDCS on neural tissue is rudimental.
Associative memory relies on a wide network, consisting of medial
temporal lobe structures and fronto-tempo-parietal network. Within
that network, the hippocampus seems to be a central hub that performs
the binding of information from different processing modules, thus
creating coherent associative memories (Davachi, 2006).
Since memory formation and retention are achieved through high
interconnectivity between hippocampal and anterior and posterior
cortical regions, many previous studies applied tDCS over frontal areas.
This line of research resulted in mixed findings. A review by Kim et al

⁎
Corresponding author at: Institute for Medical Research, Dr Subotića 4, PO Box 124, Belgrade, Serbia.
E-mail address: jovana.bjekic@imi.bg.ac.rs (J. Bjekić).

https://doi.org/10.1016/j.nlm.2018.12.007
Received 24 July 2018; Received in revised form 9 October 2018; Accepted 12 December 2018
Available online 13 December 2018
1074-7427/ © 2018 Elsevier Inc. All rights reserved.
J. Bjekić et al.
Fig. 1. Procedure and associative memory tasks. Description: A – Design of experimental session; B – Face-word associative memory task; and C – Object-location
memory task.
suggested that frontal tDCS may be effective only if the stimulation is
applied during encoding (Kim, Ekstrom, & Tandon, 2016). One recent
study showed evidence of improved face-name associative memory
following anodal stimulation of dorsolateral prefrontal cortex, in
younger but not in older adults (Leach, McCurdy, Trumbo, Matzen, &
Leshikar, 2018). Two studies found the effect on recall but no effects on
recognition of face-name pairs (Leshikar et al., 2017; Matzen, Trumbo,
Leach, & Leshikar, 2015), while Leach et al (Leach, McCurdy, Trumbo,
Matzen, & Leshikar, 2016) reported no effect on recall and negative
effect on recognition in older adults. Moreover, no effects in offline
tDCS protocol over frontal lobe were found for episodic (Medvedeva
et al., 2018) and verbal associative memory (de Lara, Knechtges,
Paulus, & Antal, 2017).
In contrast, studies that targeted posterior brain regions showed
more promising results. One research showed that tDCS over lateral
posterior parietal cortex (PPC), but not prefrontal cortex, decreases
false recognition, as well as memory bias in item and source dis-
crimination (Pergolizzi & Chua, 2016). Moreover, tempo-parietal tDCS
improved memory of nonword-picture pairs (Perceval, Martin,
Copland, Laine, & Meinzer, 2017) and object-location association in
older adults (Floel et al., 2012). Finally, a rTMS study showed that
multiple-day stimulation of PPC has immediate and long-term effects on
associative memory, due to the high connectivity with hippocampus
(Wang et al., 2014). Still, the research on the effects of tDCS over PPC
on associative memory is very limited.
Here we present two experiments in which we explore the effects of
anodal tDCS over PPC on the most frequently employed behavioral
associative memory paradigms. That is, following a network approach
to modulating memory (Kim et al., 2016), we expect that by stimulating
one node (i.e. PPC) of the broader network that the other cortical
memory hubs within the network will be activated, thus enabling
within-network synchronization that will lead to better memory per-
formance. In the first experiment we tested the effect of tDCS over left
PPC on face-word associative memory, and in the second, the effect of
tDCS over right PPC on object-location memory. We hypothesize that
anodal stimulation of parietal areas would lead to higher performance
in both associative memory tasks due to high connectivity between PPC
and hippocampus (Wang et al., 2014).
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Neurobiology of Learning and Memory 157 (2019) 114–120
2. Material and methods
2.1. Participants
Forty-two healthy participants aged 20–35 years were randomly
assigned to either first or second experiment. One participant dropped
out, which resulted in 20 participants (11 females, mean ± SD age:
26.40 ± 3.71) taking part in Experiment 1 and another 21 (12 females,
mean ± SD age: 24.15 ± 2.74) in Experiment 2. The sample size was
calculated a priori for the expected effect size (η2) of 0.15 and power
(1 − β error probability) of 0.95. All participants were naïve to tDCS,
right-handed (Edinburgh Handedness Inventory (Veale, 2014) laterality
quotient of at least 80), had normal or corrected-to-normal vision, and
reported no history of migraines, head injuries, seizures, epilepsy and
other neurological or psychiatric disorders. At the time of the experi-
ment, none of the participants was pregnant or taking medication
(apart from contraceptive pill), and was showing no signs of emotional
distress as measured by Depression Anxiety Stress Scale (Lovibond &
Lovibond, 1995), i.e. scores were between 0–9 for Depression, 0–7 for
Anxiety and 0–14 for Stress. The study was conducted in accordance
with the latest version of the Declaration of Helsinki and was approved
by the Institutional Ethics Board. All participants signed informed
consent to participate and were thoroughly debriefed after completing
the study.
2.2. Design and procedure
The study adopted double-blind, cross-over sham-controlled design,
with stimulation condition (anodal/sham) and trial (first/second) as
within-subject factors. We opted for within-subjects design in order to
control for individual differences in current responsiveness and cogni-
tive abilities (Li, Uehara, & Hanakawa, 2015). Within both experiments,
the order of stimulation conditions was counterbalanced across parti-
cipants and parallel forms of associative memory tasks were used to
prevent the possible confounding effect of learning (Thair, Holloway,
Newport, & Smith, 2017). Experimental sessions were separated 7 days
to minimize carryover effects (Biabani, Farrell, Zoghi, Egan, &
Jaberzadeh, 2017). The experiments were carried out between 10 am
and 4 pm, however, it was arranged for each participant to have both
J. Bjekić et al.
experimental sessions at the same time of the day to reduce the risk of
circadian influences (Li et al., 2015).
In each experimental session (Fig. 1A), participants first underwent
20-minute tDCS treatment (either anodal or sham), during which they
played a low-demand noncompetitive computer game. The rationale
behind introducing the game was to control for possible interference
effects of unstructured activities performed during tDCS (Horvath,
Carter, & Forte, 2014). In Experiment 1 participants played match-pairs
memory game, and in Experiment 2 they played a tile-matching puzzle
game. In match-pairs game participants need to match pairs of tiles
with the same pattern, by selecting two tiles in each turn. In tile-
matching puzzle, participants had to eliminate the balls rolling around
the screen by targeting them with a color-matched ball from player’s
deck. In spite of their differences, both tasks required maintaining
constant level of attention and manipulation of information in short-
term memory storage. The selection of the tasks was based on the as-
sumption that minimal engagement of same cortical circuits during
stimulation may promote tDCS effects (Nozari, Woodard, & Thompson-
Schill, 2014). Different games were chosen for Experiments 1 and 2 in
order to avoid proactive interference effects on the associative memory
task. In both games, participants started at the beginner’s level (e.g.
3 × 4 tiles in match-pair memory game), and progressed to upper levels
each time reached an accuracy of 85%. This way, the difficulty was
individually-calibrated to occupy attention, but not to exhaust partici-
pants. Furthermore, every 5 min during tDCS (i.e. at minutes 3, 8, 13,
and 18) participants were asked to rate the unpleasantness of the
treatment on a 10-point scale (1 – highly comfortable; 10 – extremely
unpleasant). In Experiment 1, following tDCS participants were pre-
sented with the face-word memory task, which consisted of the practice
session and two trials. In the practice session, participants were pro-
vided with detailed instruction and demonstration of the task, alongside
a few examples. Trial 1 and trail 2 of the face-word memory task were
identical, each comprising of learning and recall block, with the same
20 target face-word pairs. Between the two trials, participants com-
pleted a verbal fluency task. This control task was introduced in order
to assess if the tDCS effects are stimulation site- and function-specific.
Experiment 2 followed the same procedure, with the only difference
being the associative memory task that assessed object location
memory. After completing both anodal and sham condition, at the end
of the second session of the respective experiment participants were
asked to guess the session in which they received real stimulation in
order to evaluate if the blinding was successful.
2.2.1. Face-word memory task
The task was designed in OpenSesame (Mathot, Schreij, &
Theeuwes, 2012), and presented on 23″ monitor (resolution:
1920 × 1080, refresh rate 60 Hz). The task consisted of learning and
recall block. In the learning block, i.e. encoding phase, 20 face-word
pairs were presented for 3000 ms each in a consecutive manner. The
faces were extracted from the FEI database (Thomaz & Giraldi, 2010)
and were all gray-scale frontal face images (360 × 260 pixels). The
words were high-frequency and high-imaginability consisting of 2 or 3
syllables (e.g. banana /banana/, prozor /window/, balon /balloon/,
etc.). In the recall block, images from the first block were mixed with 30
new distractor faces and these 50 images were consecutively presented
in a pre-randomized order. Participants were first asked to identify if
the face was presented in the learning block and if so to recall the word
that was presented alongside it (Fig. 1B). Thus, in this block partici-
pants had first to perform item-memory task, and then proceed to as-
sociative memory free recall. The number of correctly recalled words
that matched correctly recognized faces was used as a measure of as-
sociative memory. Parallel forms of the task were designed by replacing
male faces with females, and vice versa, and matching words in terms of
frequency, imaginability, length, and category (e.g. breskva /peach/ in
one form and višnja /cherry/ in the other).
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Neurobiology of Learning and Memory 157 (2019) 114–120
2.2.2. Object-location memory task
The same software and equipment were used for the design and
administration of stimuli as in face-memory task. The object-location
task consisted of learning and recall block, as well. In the learning
block, participants were presented with color images of 15 everyday
objects consecutively appearing in a 4 × 4 grid (16 positions in total)
for 1000 ms each. The location of the object followed pseudo-random
pattern i.e. seven positions were unique (1 object per position), four
positions were shared (2 objects per position) and no object appeared at
five positions. The objects were highly familiar, i.e. met frequently in
everyday life (e.g. chair, spoon) or represented typical examples of
common categories (e.g. animals, musical instruments, vehicles). In the
recall block participants were presented with an image of each object in
pre-randomized order and were asked to click on its location. Thus, for
each object, there was a 6% chance of random guessing. In this task, the
measure of associative memory was the number of correctly matched
objects and within-grid locations. Parallel forms of the task were de-
signed by 90-degrees rotation of the positions within the grid and
matching objects for category and color (e.g. apple in one form and
strawberry in the other).
2.2.3. Verbal fluency task
In the verbal fluency task, participants were presented with an al-
phabet letter and instructed to list as many unique words starting with
the given letter. The time limit for this task was set to 45 s. This task is
widely used to access frontal lobe functions such as executive control
functions (Shao, Janse, Visser, & Meyer, 2014). The letters used in the
fluency test (R and M) were randomized across sessions and stimulation
conditions.
2.3. Transcranial direct current stimulation
The procedure followed contemporary safety standards and guide-
lines of tDCS application in human studies (Bikson et al., 2016). The
tDCS was delivered through constant current stimulator (Jonos-4,
Electronic Design Medical D.O.O., Belgrade, Serbia) via rubber elec-
trodes inserted in saline-soaked sponges positioned over PPC (anodal
electrode, size: 5 × 5 cm) and contralateral cheek (reference electrode,
size: 5 × 5 cm). The anodal electrode was placed over left PPC in Ex-
periment 1 (P3 10-20 EEG system) and right PPC in Experiment 2 (P4
10-20 EEG system) (Fig. 2A). In the anodal condition, the participants
were administrated 1.5 mA (0.06 mA/cm2) of constant current for
20 min (15 s fade in/out), while in the sham condition the current was
administrated during first and last 30 s of the treatment period. The
simulation of local electric fields generated by tDCS, that was per-
formed in Comets software (Jung, Kim, & Im, 2013), showed that this
montage covers respective PPC areas (Fig. 2B). This particular electrode
placement is chosen in order to target wide areas of neocortical memory
network with anodal electrode, thus enhancing excitability in the par-
ietal hippocampal network, and avoid any potential inhibitory effect
under cathodal electrode as it was placed extracranially.
2.4. Data analysis
Paired-samples t-test was used to asses differences between anodal
and sham condition with regard to the unpleasantness of the tDCS
procedure and the control task. Statistical significance for main effects
and interaction was determined using 2 × 2 repeated measures ANOVA
with stimulation condition (anodal/sham) and trial (first/second) as
within-subject factors; and the associative memory performance as
dependent variable, i.e. the number of correctly recalled face/word
pairs in Experiment 1 and the number of correctly recalled object-lo-
cation pairs in Experiment 2. Moreover, MANOVAs, with follow-up
ANOVAs were used to test the effect of possible confounding factors, i.e.
participants’ gender (male/female), stimulation order (1st/2nd ses-
sion), task form (A/B) on memory performance across trials and
J. Bjekić et al. Neurobiology of Learning and Memory 157 (2019) 114–120

Fig. 2. tDCS montage. Description: A – Electrode placement in Experiment 1 (anode, marked red, over left PPC) and Experiment 2 (anode over right PPC); B –
Simulation of electric fields for 1.5 mA with PPC-contralateral cheek montage of tDCS (1 – anode; 2 – cathode), where red indicates positive electric potential.

conditions. Finally, the tDCS unpleasantness ratings were correlated Table 1

(Spearman’s rho) with associative memory performance for each con- Task performance (mean scores and standard deviations) under anodal and
dition and trial. sham stimulation in both trials.
Anodal stimulation Sham stimulation
3. Results
Trial 1 Trial 2 Trial 1 Trial 2
3.1. tDCS procedure evaluation
Experiment 1 Face-word task 5.00 10.00 3.80 8.20
(2.88) (3.32) (2.19) (3.08)
Overall, participants found the procedure only moderately un- Fluency task 13.30 (3.95) 13.15 (3.36)
pleasant and rated it below 5 for all time points on 1–10 Likert type Experiment 2 Object-location 7.48 8.48 6.24 7.48
scale. The unpleasantness ratings were slightly higher for anodal than task (2.99) (3.60) (2.63) (3.40)
sham condition [Experiment 1: Manodal = 2.63, SD = 1.90; Fluency task 11.71 (2.86) 12.29 (2.74)
Msham = 2.02, SD = 1.43; t (19) = 2.221, p = .039; Experiment 2:
Manodal = 3.16, SD = 2.01; Msham = 2.04, SD = 1.04; t (20) = 2.522,
p = .020]. Still, the guessing of stimulation conditions was not above between two trials, but stimulation had no effect on learning slope, i.e.
the chance level (40% of accuracy in Experiment 1; 38% of accuracy in the difference in performance between two trials did not differ between
Experiment 2), thus confirming the successful blinding of the tDCS anodal and sham condition.
procedure. The results of the Experiment 2, followed the same pattern. The
repeated measures ANOVA with object-location recall as output vari-
3.2. Effects on associative memory able showed the effect of stimulation condition [F(1, 20) = 4.516,
p = .046, η2 = 0.184], as well as the effect of learning [F(1,
Descriptive statistics (means and standard deviations) for all out- 20) = 7.160, p = .015, η2 = 0.264], but again without interaction ef-
come measures, i.e. associative memory performance and verbal flu- fect [F(1, 20) = 0.100, p = .755].
ency are presented in Table 1.
In the Experiment 1, the repeated measures ANOVA showed the 3.3. Control task and possible confounding effects
effect of stimulation condition [F(1, 19) = 5.446, p = .031,
η2 = 0.223], as well as the effect of learning [F(1, 19) = 110.16, No effect of tDCS was observed on verbal fluency task in Experiment
p < .001, η2 = 0.853], but no interaction effect [F(1, 19) = 0.872, 1(t (19) = 0.212, p = .843), nor in Experiment 2 (t (20) = 0.890,
p = .362]. In other words, face-word memory performance was higher p = .384). In other words, the performance on control task was not
after anodal stimulation, participants showed an increase in recall modulated by tDCS in neither of the experiments.

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J. Bjekić et al.
There were no confounding effects on memory performance in ei-
ther of the experiments. The analysis showed no effect of participants'
gender [Experiment 1: F(4, 9) = 0.974, p = .467; Experiment 2: F(4,
10) = 2.557, p = .104], no effect of stimulation order [Experiment 1: F
(4, 9) = 0.371, p = .824; Experiment 2: F(4, 10) = 0.377, p = .820],
and no effect of task form [Experiment 1: F(4, 15) = 1.172, p = .385;
Experiment 2: F(4, 10) = 0.410, p = .798.]. Moreover, the tDCS un-
pleasantness ratings did not correlate with associative memory perfor-
mance in any of the conditions (p > .500 for both trials within both
experiments).
4. Discussion
This study investigated the effects of anodal tDCS over PPC on as-
sociative memory in two separate yet closely related experiments. In
order to explore the robustness of the effects, the first experiment as-
sessed face-word associative memory, while the second tested the ob-
ject-location memory performance. In both experiments, the associative
memory performance increased following a single session of anodal
tDCS over PPC, while performance on control task remained the same.
Thus, the current study demonstrates the potential of posterior stimu-
lation site and challenges the notion that single tDCS session has no
reliable effect on memory performance (Horvath, Forte, & Carter, 2015;
Price & Hamilton, 2015). It could be argued that the inconsistencies in
findings of previous studies may stem from the choice of stimulation
site (anterior vs posterior regions), and other stimulation parameters, as
well as the heterogeneous outcome measures used to assess relational/
associative memory.
Even though face-word and object-location tasks are the two most
frequently employed associative memory paradigms, and often used
interchangeably, they differ in several important aspects. On one hand,
face-word memory relies on both visual and verbal encoding of in-
formation and requires cross-modal binding. On the other hand, object-
location memory is expected to engage visuospatial processing and
intra-modal binding. Although the underlining processes involved in
cross-modal and intra-modal binding are still a matter of debate (Del
Gatto, Brunetti, & Delogu, 2016), there is a great deal of evidence that
retrieval of different types of materials is associated with distinct brain
regions (Galli & Otten, 2011). Namely, neuroimaging studies have
shown that the memory of words primarily activates frontal and tem-
poral areas of the left hemisphere, whereas memory of objects and faces
activates posterior parts of the right hemisphere (Kim et al., 1999;
Simons, Graham, Owen, Patterson, & Hodges, 2001). Still, fMRI studies
provide evidence for parietal old/new effect in recollection of both
verbal and visual memories (Guerin & Miller, 2009). Furthermore, an
EEG-fMRI study showed that the amplitude of parietal recollection ef-
fect is associated with the activation of posterior hippocampus, para-
hippocampal cortex and retrosplenial cortex for different types of sti-
muli (Hoppstädter, Baeuchl, Diener, Flor, & Meyer, 2015). Finally, a
large body of evidence speaks to the central role of hippocampus and
adjacent structures in constituting relational memories across mod-
alities (Horecka et al., 2018; Konkel & Cohen, 2009; Olsen, Moses,
Riggs, & Ryan, 2012). Overall, these findings suggest that parietal
cortex may be the most appropriate target for neurostimulation aiming
to enhance memory performance in different modalities due to its high
functional connectivity with hippocampus as a key relay for cross-
modal binding and memory formation.
Unlike some previous studies (de Lara et al., 2017; Leach et al.,
2016, 2018; Leshikar et al., 2017; Matzen et al., 2015; Medvedeva
et al., 2018), the montage of electrodes in present experiments targeted
areas of posterior cortex which have been shown to play important role
in formation and retrieval of relational memories. Hence the positive
effects of anodal stimulation over PPC could be attributed to the fa-
cilitation of modality-general binding process and subsequent retrieval
of relational memories within the hippocampal-parietal circuit. The
consistent absence of the effect on verbal fluency supports this
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Neurobiology of Learning and Memory 157 (2019) 114–120
conclusion and further implies that the observed effects are stimulation
site- and task-specific. Namely, it seems that anodal stimulation over
PPC affects processes that are engaged in associative memory without
affecting other cognitive processes. In addition, the fact that the similar
effect sizes are obtained in both face-word and object-location experi-
ment adds to the robustness of the parietal tDCS effects and supports
the notion that posterior regions are highly engaged in multimodal
associative memory. Lastly, given that there were no effects of stimu-
lation order, task form or participants’ gender, as well as no correlation
between unpleasantness and memory performance, we can exclude that
any of these factors mediated the tDCS effect on associative memory in
either of the experiments.
Another difference between previous studies and this one is the
game played during tDCS. Namely, in both experiments participants
performed a cognitively engaging game with non-interfering material
(e.g. color patterns and face-word pairs). There are indications that
engaging target cortical circuits during stimulation may promote the
after-effects of tDCS (Nozari et al., 2014). It could be argued that cog-
nitive activity during tDCS contributed, at least partially, to the sub-
sequent effects on associative memory.
As expected, the significant effect of between-trials learning was
observed – i.e. the participants showed a positive learning curve, thus
the performance in the second trial was consistently higher than in the
first. But surprisingly, tDCS had no effect on learning slope – the
amount of new information encoded during the repeated presentation
of the stimuli (i.e. learning block) was equal across conditions. Since
this finding was consistent across both experiments, it cannot be at-
tributed to properties of the task, such as material or relative item-
difficulty. Furthermore, other studies that varied the number of
learning trials also showed no interaction of stimulation condition
(anodal/sham) and the number of stimuli presentations (once/twice)
(Leshikar et al., 2017). One possible explanation is that the tDCS pre-
dominantly modulated the activity of hippocampal subregions that
underline retrieval, but not encoding of previously formed associations.
This functional subdivision of hippocampus during encoding and re-
trieval of relational memories was shown in high-resolution fMRI study
(Zeineh, Engel, Thompson, & Bookheimer, 2003), but the behavioral
distinction of these processes is very difficult to demonstrate in any
cognitive task. The only possible experimental dissociation would have
to rely on the assumption that multiple presentations of stimuli enhance
encoding but not the recall component of memory performance. That is,
repeated presentations enable better encoding, while the retrieval of
the information is only indirectly affected, as the deeper encoded in-
formation is easier to retrieve. Thus, the increased performance fol-
lowing multiple presentation trials does not per se reveal the process
that has been facilitated. Still, the difference or absence of differences
between sham and active stimulation condition in the learning slope
could indicate if tDCS affected both encoding and recall or just recall,
since the number of presentations was the same in both conditions.
Bearing all this in mind, we could tentatively argue that tDCS dom-
inantly facilitated recall of relational memories. In order to provide
more compelling evidence further studies that would combine high-
definition neuroimaging techniques and neurostimulation are needed.
We accounted for the type of material to be retrieved by placing the
active electrode over left PPC for the recall of verbal stimuli and over
right for the recall of visuospatial information. The lateralization of
visual and verbal material processing has been well documented (Kelley
et al., 1998; Weber, Fliessbach, Lange, Kügler, & Elger, 2007), espe-
cially in neuropsychological studies (Cohen, 1992; Wagner, Sziklas,
Garver, & Jones-Gotman, 2009). Our study shows that unilateral tDCS
affects associative memory performance in a material congruent
manner. Still, some studies, challenge traditional notions on functional
lateralization and provide evidence for bilateral activation while pro-
cessing verbal, as well as visual material (e.g. (Guerin & Miller, 2009;
Rugg & Vilberg, 2013). Further studies are needed to test the re-
lationship between left/right stimulation site and the type of associative
J. Bjekić et al.
memory material.
5. Conclusion
A single-session of anodal tDCS over posterior parietal cortex can
induce function-specific changes in associative memory performance in
both face-word and object-location task. This study produced evidence
in favor of parietal cortex as a promising stimulation target for memory
enhancement.
Acknowledgments
This work was supported by a Grant [OI175012] from the Ministry
of Education, Science and Technological Development of Republic of
Serbia. The authors would like to thank Jelena Vujičić (Experiment 1)
and Milena Dimitrijević (Experiment 2) for technical support and as-
sistance in data collection.
Declaration of interest
Authors confirm that there are no known conflicts of interest asso-
ciated with this publication and there has been no significant financial
support for this work that could have influenced its outcome.
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