Research Article

Effects of Transcranial Alternating Current Stimulation on Cognitive Functions in Healthy Young and Older Adults

Table 1

Summary of studies that have applied tACS to enhance cognitive functions in healthy adults, in alphabetical order.

AuthorsParticipants and designDomain, task, and DVElectrode montage Stimulation parametersMain resultsConclusion

Ambrus et al. 2015 [45] (age )
Crossover design (tACS versus sham)
Memory/declarative memory
Associative word-pair learning test
Accuracy (cued recall) after encoding and after a night of sleep
L DLPFC (F3) + L mastoid/R DLPFC (F4) + R mastoidFrequency: 140 Hz
Intensity: 1 mA peak-to-peak amplitude 
Duration: 10 min
Electrode size: 5 × 5 cm2 (F3/4) and 5 × 7 cm2 (mastoids)
Timing: task (encoding)
Forgetting (accuracy decrease) after a night of sleep in sham but not tACS condition
#Offline-effects
Potential effect of 140 Hz-tACS on consolidation

Hoy et al. 2015 [34] (age )
Crossover design (tACS versus tDCS versus sham)
Executive function/working memory
2- and 3-back task before and after stimulation
Change in -primes and accurate RT
F3 + R supraorbitalFrequency: 40 Hz (tACS); NA (tDCS)
Intensity: 1.5 mA peak-to-peak amplitude (tACS); 2 mA (tDCS)
Duration: 20 min
Electrode size: both 5 × 7 cm2
Timing: task (2-back)
Selective improvement on 3-back after gamma-tACS compared to tDCS and sham
No effects of RT
#Offline-effects
Role of frontal gamma-tACS in neuromodulation
Preliminarily evidence for gamma-tACS preferentially improving working memory performance at higher loads

N. Jaušovec and K. Jaušovec 2014 [35] in P3 and in F3 tACS group (age )
Crossover within-group design (tACS versus sham)
Attention/working memory
Visual-array comparison task
EEG ERP amplitudes and latencies and memory capacity score
P3 (L) or F3 (L) + R supraorbital (Fp2)Frequency: 5 ± 1 Hz (based on individual alpha peak minus 5 Hz)
Intensity: 1–2.2 mA (based on individual skin sensations)
Duration: 15 min
Electrode size: both 5 × 7 cm2
Timing: rest
Increased capacity score after P3-tACS and no effect after F3-tACS
Parietooccipital P300 ERP component latency decrease
#Offline-effects
Causal relation between working memory storage capacity and theta frequency oscillations in the left parietal area

Jaušovec et al. 2014 [36] in P3 and in P4 and in F3 tACS group (age )
Crossover within-group design (tACS versus sham)
Executive function/working memory
Corsi-block tapping, digit span (both forward and backward), and -back
Span core and number of correct responses
P3 (L), P4 (R), or F3 (L) + R supraorbital (Fp2)Frequency: 5 ± 1 Hz (based on individual alpha peak minus 5 Hz)
Intensity: 1–2.2 mA (based on individual skin sensations)
Duration: 15 min
Electrode size: both 5 × 7 cm2
Timing: rest
Increased capacity score after L and R parietal tACS, but not frontal, compared to sham
More pronounced effect of L parietal tACS on backward compared to forward recall
#Offline-effects
Central role of parietal areas for working memory storage capacity

Meiron and Lavidor 2014 [37] in tACS and in sham (age 19–27)
Between-subjects design
Executive function/working memory
Verbal working memory task (2-back task with word pairs) and rating of success-confidence on task
Accuracy and self-evaluation scores
L DLPFC (F3/AF3 midpoint) + R DLPFC (F4/AF4 midpoint)Frequency: 4.5 Hz
Intensity: .5 mA peak-to-peak amplitude 
Duration: 20 min
Electrode size: 4 × 4 cm2
Timing: task (verbal -back)
Improved online accuracy in bilateral tACS compared to sham
WM enhancement related to self-evaluation scores
#Online-effects
Increased functional connection between working and retrospective monitoring through tACS

Pahor and Jaušovec 2014 [20] in parietal and in frontal tACS group (age )
Crossover within-group design (tACS versus sham)
Executive function/cognitive control/fluid intelligence (problem-solving and visual-spatial reasoning)
Progressive matrices, paper folding, and cutting test
EEG frequency spectra and performance test scores
L parietal (P3) or L frontal (F3) + R supraorbital (Fp2)Frequency: 5 ± 1 Hz (based on individual alpha peak minus 5 Hz)
Intensity: 1–2.2 mA (based on individual skin sensations)
Duration: 15 min
Electrode size: 5 × 7 cm2 (P3/F3) and 7 × 10 cm2 (Fp2)
Timing: rest
Increased resting EEG theta power and decreased EEG alpha power after theta-tACS
Improved test scores after tACS
Behavioral effect more pronounced in L parietal tACS group (better in both tests) compared to L frontal
#Offline-effects
Influence of brain oscillatory activity by theta-tACS
Importance of parietal regions for fluid intelligence
Parietal tACS increased working memory storage capacity and frontal tACS influenced attentional components

Polanía et al. 2012 [38] and in control experiment (age 22–30)
Crossover design (tACS versus sham)
Executive function/working memory
Delayed letter discrimination task
RT and correct responses
L frontal (F3) + L parietal (P3)
Cz (reference electrode)
Frequency: 6 Hz (35 Hz in control experiment)
Intensity: 1 mA peak-to-peak amplitude 
Duration: 14 min
Electrode size: all 5 × 5 cm2
Timing: task (letter discrimination)
Reduced RT with synchronized 6 Hz-tACS
Increased RT with desynchronized 6 Hz-tACS
#Online-effects
Causality of frontoparietal theta phase coupling
Suitability of tACS to induce coupling and decoupling of behaviorally relevant brain rhythms

Polanía et al. 2015 [43] = 86 (age 20–30)
Crossover design (alternating intervals of stimulation/no stimulation)
Executive function/decision-making
Perceptual- and value-based decisions
RT and accuracy
mPFC (Fpz) + parietal (Pz)
R shoulder (reference electrode)
Frequency: 55 Hz (modulated by a 6 Hz envelope)
Intensity: 2 mA peak-to-peak amplitude 
Duration: ~18 min
Electrode size: 5 × 7 cm2 (Fpz/Pz) and 10 × 10 cm2 (shoulder)
Timing: task (decision-making)
More inaccurate value-based decisions with desynchronized tACS
No effect on perceptual decisions
#Online-effects
Causal influence of degree of rhythmic synchronization between frontoparietal brain areas on value-based decision-making

Santarnecchi et al. 2013 [39] (age )
Crossover design (5 conditions: sham, theta-, alpha-, beta-, and gamma-tACS)
Executive function/cognitive control/fluid intelligence (problem-solving and visual-spatial reasoning)
Progressive matrices
RT and accuracy
L MFG (−34, 16, 30) + CzFrequency: 5, 10, 20, or 40 Hz
Intensity: .75 mA peak-to-peak amplitude 
Duration: delivered for duration of task performance
Electrode size: 5 × 7 cm2
Timing: task (visuospatial reasoning)
Shortening of RT in gamma-tACS condition
Selective enhancement of more complex trials
#Online-effects
Causal involvement of gamma band synchronization in higher-order cognition

Santarnecchi et al. 2016 [44] (age ) in Exp. 1 and (age ) in Exp. 2
Crossover within-group design (3 conditions in Exp. 1: sham, theta-, and gamma-tACS; 3 conditions in Exp. 2: sham, tRNS, and gamma)
Executive function/cognitive control/fluid intelligence and working memory
Abstract-reasoning task (progressive matrices), delayed-match-to-sample (change-localization) task, and control task (odd-even numbers)
RT and accuracy
L MFG (−34, 16, and 30) + CzFrequency: 5, 40 Hz (Exp. 1); 5, 101–640 Hz tRNS (Exp. 2)
Intensity: .75 mA peak-to-peak amplitude 
Duration: 30 min
Electrode size: 5 × 5 cm2
Timing: task (visuospatial reasoning and working memory)
Shortening of RT to solve complex logic problem in gamma-tACS condition compared to all other conditions in both Exp. 1 and Exp. 2
Greater tACS-induced enhancement for slower participants
No effect in theta-tACS or tRNS
No effect on working memory task
#Online-effects
Frequency-specific neuromodulatory effects on cognitive ability
Individual differences in gamma-tACS responsiveness

Sela et al. 2012 [40] in L, in R, and in sham
Between-subjects design (3 groups: L, R, and sham)
Executive function/cognitive control/decision-making
Balloon Analog Risk Task
Number of pumps and total number of balloon explosions
L DLPFC (F3) + L temporal (CP5)/R DLPFC (F4) + R temporal (CP6)Frequency: 6.5 Hz
Intensity: 1 mA peak-to-peak amplitude 
Duration: 15 min
Electrode size: all 5 × 5 cm2
Timing: task (decision-making)
Riskier decision-making in L-tACS group compared to R-tACS and sham group
No difference between sham and R-tACS groups
#Online-effects
Theta-band oscillatory and DLPFC activity critical for decision-making
Disruption of ability to process and adjust actions based on negative feedback and errors by theta-tACS

van Driel et al. 2015 [41] (age 19–31)
Crossover design (theta- versus alpha-tACS)
Executive function/cognitive control/conflict processing
Spatial response conflict task (color-location Simon task)
RT and accuracy
Centered between FCz/Cz bilaterally on cheeks (reference electrodes)Frequency: 8–12 versus 4–8 Hz (based on individual alpha and theta peak)
Intensity: 2 mA peak-to-peak amplitude 
Duration: ~20 min
Electrode size: 9 cm2 (FCz/Cz) and 35 cm2 (cheeks)
Timing: task (spatial conflict)
Slower RTs and lower accuracy for high conflict trials: conflict reduced during tACS
Effect driven by slower RTs in low-conflict trials
#Online-effects
Modulation of cognitive control through theta-tACS

Vosskuhl et al. 2015 [42] in tACS and in sham group (age )
Between-subjects design
Executive function/working memory (WM) and short-term memory (STM)
Digit span backward and 3-back task (both WM) and digit span forward (STM)
EEG frequency spectra, mean list length (digit span task), and accuracy (3-back task)
FPz + PzFrequency: ~3.7–4.6 Hz (below individual theta frequency)
Intensity: ~.4–1.3 mA (below individual phosphene or perception threshold)
Duration: 18 min
Electrode size: all 5 × 7 cm2
Timing: task (digit span)
Increased digit span forward during stimulation in tACS group
No effect in digit span backward and 3-back accuracy
No effect beyond the stimulation period
Increase in theta-amplitude after stimulation in tACS group
#Online- + offline-effects
Successful manipulation of theta frequencies caused increase in STM capacity

EEG, electroencephalography; ERP, event-related potential; DLPFC, dorsolateral prefrontal cortex; DV, dependent variable; L, left; MFG, middle frontal gyrus; mPFC, medial prefrontal cortex; R, right; RT, reaction times; tACS, transcranial alternating current stimulation; tDCS, transcranial direct current stimulation.