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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Psychol.</journal-id>
<journal-title>Frontiers in Psychology</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Psychol.</abbrev-journal-title>
<issn pub-type="epub">1664-1078</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fpsyg.2024.1479887</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Psychology</subject>
<subj-group>
<subject>Mini Review</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Neuromodulation on the ground and in the clouds: a mini review of transcranial direct current stimulation for altering performance in interactive driving and flight simulators</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name><surname>Sansevere</surname> <given-names>Kayla S.</given-names></name>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/2815738/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/investigation/"/>
<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
<role content-type="https://credit.niso.org/contributor-roles/project-administration/"/>
<role content-type="https://credit.niso.org/contributor-roles/validation/"/>
<role content-type="https://credit.niso.org/contributor-roles/visualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Ward</surname> <given-names>Nate</given-names></name>
<uri xlink:href="https://loop.frontiersin.org/people/140964/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/project-administration/"/>
<role content-type="https://credit.niso.org/contributor-roles/supervision/"/>
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</contrib>
</contrib-group>
<aff><institution>Tufts Applied Cognition Laboratory, Department of Psychology, Tufts University</institution>, <addr-line>Medford, MA</addr-line>, <country>United States</country></aff>
<author-notes>
<fn fn-type="edited-by" id="fn0001">
<p>Edited by: Adam Joseph Toth, University of Limerick, Ireland</p>
</fn>
<fn fn-type="edited-by" id="fn0002">
<p>Reviewed by: Abhishek Datta, Soterix Medical Inc., United States</p>
</fn>
<corresp id="c001">&#x002A;Correspondence: Kayla S. Sansevere, <email>kayla.sansevere@tufts.edu</email></corresp>
</author-notes>
<pub-date pub-type="epub">
<day>28</day>
<month>11</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>15</volume>
<elocation-id>1479887</elocation-id>
<history>
<date date-type="received">
<day>13</day>
<month>08</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>18</day>
<month>11</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2024 Sansevere and Ward.</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>Sansevere and Ward</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Transcranial direct current stimulation (tDCS) has emerged as a promising tool for cognitive enhancement, especially within simulated virtual environments that provide realistic yet controlled methods for studying human behavior. This mini review synthesizes current research on the application of tDCS to improve performance in interactive driving and flight simulators. The existing literature indicates that tDCS can enhance acute performance for specific tasks, such as maintaining a safe distance from another car or executing a successful plane landing. However, the effects of tDCS may be context-dependent, indicating a need for a broader range of simulated scenarios. Various factors, including participant expertise, task difficulty, and the targeted brain region, can also influence tDCS outcomes. To further strengthen the rigor of this research area, it is essential to address and minimize different forms of research bias to achieve true generalizability. This comprehensive analysis aims to bridge the gap between theoretical understanding and practical application of neurotechnology to study the relationship between the brain and behavior, ultimately providing insights into the effectiveness of tDCS in transportation settings.</p>
</abstract>
<kwd-group>
<kwd>transcranial direct current stimulation</kwd>
<kwd>tDCS</kwd>
<kwd>driving simulator</kwd>
<kwd>flight simulator</kwd>
<kwd>transportation</kwd>
</kwd-group>
<counts>
<fig-count count="0"/>
<table-count count="2"/>
<equation-count count="0"/>
<ref-count count="96"/>
<page-count count="7"/>
<word-count count="6838"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Cognition</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="sec1">
<label>1</label>
<title>Introduction</title>
<p>There is robust interest in using non-invasive brain stimulation (NIBS) to characterize and modulate human cognition and behavior (<xref ref-type="bibr" rid="ref4">Antal et al., 2022</xref>; <xref ref-type="bibr" rid="ref9">Berryhill, 2014</xref>; <xref ref-type="bibr" rid="ref11">Bikson et al., 2018</xref>; <xref ref-type="bibr" rid="ref28">Dubljevi&#x0107; et al., 2014</xref>; <xref ref-type="bibr" rid="ref93">Wexler, 2017</xref>, <xref ref-type="bibr" rid="ref94">2022</xref>). NIBS methods have potential applications in demanding scenarios that require cognitive, perceptual, and motor skills, such as driving a car or piloting an aircraft. Both drivers and pilots often encounter challenging situations, like making quick decisions at busy intersections or landing in adverse weather conditions. Performance can also suffer in monotonous situations, such as during partially automated driving (<xref ref-type="bibr" rid="ref59">McWilliams and Ward, 2021</xref>).</p>
<p>Transcranial direct current stimulation (tDCS) is a prominent NIBS technique that delivers weak electrical currents via scalp electrodes to initiate subthreshold membrane polarization and alter neuronal activity (<xref ref-type="bibr" rid="ref65">Nitsche et al., 2008</xref>; <xref ref-type="bibr" rid="ref66">Nitsche and Paulus, 2000</xref>, <xref ref-type="bibr" rid="ref67">2001</xref>). The electrical current flows into the brain through the anode, which likely increases cortical excitability through depolarization, and exits through the cathode, reducing excitability via hyperpolarization (<xref ref-type="bibr" rid="ref52">Liu et al., 2018</xref>). If a specific brain region is involved in a task, modulating neuronal excitability could enhance or inhibit performance on that task (<xref ref-type="bibr" rid="ref45">Knotkova et al., 2019</xref>). Typical current intensities range from 1 to 2.5&#x202F;mA, although currents as high as 4&#x202F;mA may be used (<xref ref-type="bibr" rid="ref20">Chhatbar et al., 2017</xref>; <xref ref-type="bibr" rid="ref43">Khadka et al., 2020</xref>; <xref ref-type="bibr" rid="ref77">Reckow et al., 2018</xref>). Stimulation should be applied for no more than 1&#x202F;h during a task (online) or before it (offline) (<xref ref-type="bibr" rid="ref96">Woods et al., 2016</xref>). While online stimulation may be ideal for immediate performance enhancement, the effects of offline tDCS can last after the stimulation ends and may be better suited for investigating longer-term neural changes (<xref ref-type="bibr" rid="ref12">Bikson and Rahman, 2013</xref>; <xref ref-type="bibr" rid="ref57">Martin et al., 2014</xref>; <xref ref-type="bibr" rid="ref61">Miniussi et al., 2013</xref>; <xref ref-type="bibr" rid="ref69">Ohn et al., 2008</xref>; <xref ref-type="bibr" rid="ref87">Stagg and Nitsche, 2011</xref>).</p>
<p>Electrodes are positioned according to standardized electroencephalography (EEG) system coordinates. In conventional tDCS, two large sponge electrodes deliver a broad current across various brain regions (<xref ref-type="bibr" rid="ref48">Kuo et al., 2013</xref>). High-definition tDCS (HD-tDCS) is a significant advancement utilizing smaller electrodes arranged closely together to achieve more focal current flow than conventional tDCS (<xref ref-type="bibr" rid="ref1">Alam et al., 2016</xref>; <xref ref-type="bibr" rid="ref24">Datta et al., 2009</xref>; <xref ref-type="bibr" rid="ref90">Villamar et al., 2013</xref>). Commonly targeted brain regions include the dorsolateral prefrontal cortex (DLPFC) and the primary motor cortex (<xref ref-type="bibr" rid="ref26">Dedoncker et al., 2016</xref>; <xref ref-type="bibr" rid="ref41">Jacobson et al., 2012</xref>). The DLPFC is linked to working memory, cognitive control, and decision-making (<xref ref-type="bibr" rid="ref6">Barbey et al., 2013</xref>; <xref ref-type="bibr" rid="ref47">Krawczyk, 2002</xref>; <xref ref-type="bibr" rid="ref55">MacDonald et al., 2000</xref>), while the primary motor cortex is associated with skill acquisition and procedural learning (<xref ref-type="bibr" rid="ref42">Karni et al., 1998</xref>). Evidence from functional near-infrared spectroscopy (fNIRS) and event-based magnetoencephalography (MEG) indicates an interaction between the prefrontal cortex and primary motor cortex during critical driving maneuvers, such as accelerating and braking, particularly under varying demands (<xref ref-type="bibr" rid="ref32">Foy and Chapman, 2018</xref>; <xref ref-type="bibr" rid="ref34">Geissler et al., 2021</xref>; <xref ref-type="bibr" rid="ref91">Walshe et al., 2022</xref>).</p>
<p>Importantly, tDCS is generally well-tolerated in both healthy individuals and clinical populations (<xref ref-type="bibr" rid="ref3">Antal et al., 2017</xref>; <xref ref-type="bibr" rid="ref5">Apar&#x00ED;cio et al., 2016</xref>; <xref ref-type="bibr" rid="ref10">Bikson et al., 2016</xref>; <xref ref-type="bibr" rid="ref71">Palm et al., 2018</xref>). Compared to other NIBS methods like transcranial magnetic stimulation (TMS), tDCS is portable and adaptable for various settings, from remotely supervised clinical trials (<xref ref-type="bibr" rid="ref74">Pilloni et al., 2022</xref>) to physically demanding activities like sprint cycling (<xref ref-type="bibr" rid="ref33">Garner et al., 2021</xref>; <xref ref-type="bibr" rid="ref39">Huang et al., 2019</xref>) and military operations (<xref ref-type="bibr" rid="ref17">Bruny&#x00E9; et al., 2020</xref>; <xref ref-type="bibr" rid="ref64">Nelson and Tepe, 2015</xref>). The cognitive and perceptual enhancement effects of tDCS on operator performance and workload have been examined using computer-based tasks like the Multi-Attribute Task Battery (MATB) (<xref ref-type="bibr" rid="ref62">Nelson et al., 2016</xref>, <xref ref-type="bibr" rid="ref63">2019</xref>; <xref ref-type="bibr" rid="ref76">Rao et al., 2024</xref>), which was developed by the National Aeronautics and Space Administration (NASA) to mirror the complex responsibilities that pilots manage in flight (<xref ref-type="bibr" rid="ref83">Santiago-Espada et al., 2011</xref>). Other gamified tasks, like NeuroRacer (<xref ref-type="bibr" rid="ref38">Hsu et al., 2015</xref>) and Space Fortress (<xref ref-type="bibr" rid="ref84">Scheldrup et al., 2014</xref>), have also been used for testing.</p>
<p>Investigating tDCS in more immersive environments could further clarify its practical applications. Interactive driving (<xref ref-type="bibr" rid="ref31">Fisher et al., 2011</xref>) and flight (<xref ref-type="bibr" rid="ref2">Allerton, 2009</xref>; <xref ref-type="bibr" rid="ref36">Hays et al., 1992</xref>) simulators offer safe, controlled settings that mimic real-life demands (<xref ref-type="bibr" rid="ref79">Roberts A. P. J. et al., 2020</xref>). Interactive simulators are effective in predicting on-road driving skills (<xref ref-type="bibr" rid="ref91">Walshe et al., 2022</xref>) and supporting pilot training (<xref ref-type="bibr" rid="ref81">Ross and Gilbey, 2023</xref>). With ongoing research supporting its effectiveness, tDCS holds promise for widespread use in cognitive and motor task enhancement. Given the consistent interest in tDCS across clinical, empirical, and commercial contexts, its potential applications for performance enhancement in transportation settings are highly relevant and merit investigation.</p>
</sec>
<sec id="sec2">
<label>2</label>
<title>Current mini review</title>
<p>This review summarizes and evaluates research on the use of tDCS to modulate driver and pilot performance. We conducted searches for refereed articles on Google Scholar and PubMed using the keywords: &#x201C;driving&#x201D; OR &#x201C;flight&#x201D; AND &#x201C;transcranial direct current stimulation (tDCS).&#x201D; This search yielded nine potentially relevant publications. Our scope included studies that recruited healthy participants from nonclinical samples and used interactive driving or flight simulators. Three studies were excluded from review because they did not meet these criteria (<xref ref-type="bibr" rid="ref16">Brunnauer et al., 2018</xref>; <xref ref-type="bibr" rid="ref18">Burkhardt et al., 2023</xref>; <xref ref-type="bibr" rid="ref75">Pope et al., 2018</xref>). Ultimately, six publications met the criteria for inclusion and were reviewed (see <xref ref-type="table" rid="tab1">Tables 1</xref>, <xref ref-type="table" rid="tab2">2</xref>).</p>
<table-wrap position="float" id="tab1">
<label>Table 1</label>
<caption>
<p>Driving simulators and tDCS summary of studies.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Study</th>
<th align="left" valign="top">Design</th>
<th align="left" valign="top">Masking</th>
<th align="center" valign="top">Sample</th>
<th align="center" valign="top">Sessions</th>
<th align="center" valign="top">Current</th>
<th align="left" valign="top">Duration</th>
<th align="left" valign="top">Montage (surface area)</th>
<th align="left" valign="top">Sham</th>
<th align="left" valign="top">Key results</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle">
<xref ref-type="bibr" rid="ref7">Beeli et al. (2008)</xref>
</td>
<td align="left" valign="middle">Mixed groups</td>
<td align="left" valign="middle">Not mentioned</td>
<td align="center" valign="middle"><italic>N</italic> =&#x202F;21<break/><italic>n</italic><sub>F3</sub> =&#x202F;10<break/><italic>n</italic><sub>F4</sub> =&#x202F;11</td>
<td align="center" valign="middle">3</td>
<td align="center" valign="middle">1&#x202F;mA</td>
<td align="left" valign="middle">15&#x202F;min offline</td>
<td align="left" valign="middle">Anode: F3 or F4<break/>cathode: ipsilateral mastoid<break/>anode: ipsilateral mastoid<break/>cathode: F3 or F4<break/>(35&#x202F;cm<sup>2</sup>)</td>
<td align="left" valign="middle">None</td>
<td align="left" valign="middle">Fewer speeding violations and more headway distance from pre-stim to post-stim if anodal than cathodal</td>
</tr>
<tr>
<td align="left" valign="middle">
<xref ref-type="bibr" rid="ref82">Sakai et al. (2014)</xref>
</td>
<td align="left" valign="middle">Within groups</td>
<td align="left" valign="middle">Single-masked</td>
<td align="center" valign="middle"><italic>N</italic> =&#x202F;13</td>
<td align="center" valign="middle">3</td>
<td align="center" valign="middle">1.5&#x202F;mA</td>
<td align="left" valign="middle">20&#x202F;min<break/>online</td>
<td align="left" valign="middle">Anode: F3 and F4<break/>cathode: F4 and F3<break/>(35&#x202F;cm<sup>2</sup>)</td>
<td align="left" valign="middle">30&#x202F;s ramp up/30&#x202F;s ramp down</td>
<td align="left" valign="middle">Fewer lane deviations and more accurate headway distance when anodal than cathodal and sham</td>
</tr>
<tr>
<td align="left" valign="middle">
<xref ref-type="bibr" rid="ref29">Facchin et al. (2023)</xref>
</td>
<td align="left" valign="middle">Within groups</td>
<td align="left" valign="middle">Not mentioned</td>
<td align="center" valign="middle"><italic>N</italic> =&#x202F;27</td>
<td align="center" valign="middle">3</td>
<td align="center" valign="middle">2&#x202F;mA</td>
<td align="left" valign="middle">20&#x202F;min<break/>online</td>
<td align="left" valign="middle">Anode: FC4<break/>cathode: Fp1<break/>(35&#x202F;cm<sup>2</sup>)<break/>anode: FC4<break/>cathodes: Cp4/FT8/AF4/FCZ<break/>(6&#x202F;cm<sup>2</sup>)</td>
<td align="left" valign="middle">10&#x202F;s ramp up/10&#x202F;s ramp down</td>
<td align="left" valign="middle">Quicker foot and hand RTs when active than sham; stronger effects when HD than conventional</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<p>RTs, reaction times.</p>
</table-wrap-foot>
</table-wrap>
<table-wrap position="float" id="tab2">
<label>Table 2</label>
<caption>
<p>Flight simulators and tDCS summary of studies.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Study</th>
<th align="left" valign="top">Design</th>
<th align="left" valign="top">Masking</th>
<th align="center" valign="top">Sample</th>
<th align="center" valign="top">Sessions</th>
<th align="center" valign="top">Current</th>
<th align="left" valign="top">Duration</th>
<th align="left" valign="top">Montage (surface area)</th>
<th align="left" valign="top">Sham</th>
<th align="left" valign="top">Key results</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle">
<xref ref-type="bibr" rid="ref21">Choe et al. (2016)</xref>
</td>
<td align="left" valign="middle">Between groups</td>
<td align="left" valign="middle">Double-masked</td>
<td align="center" valign="middle"><italic>N</italic> =&#x202F;32<break/><italic>n</italic><sub>DLPFC</sub> =&#x202F;14<break/>(<italic>n</italic><sub>active DLPFC</sub> =&#x202F;7)<break/><italic>n</italic><sub>M1</sub> =&#x202F;18<break/>(<italic>n</italic><sub>active M1</sub> =&#x202F;10)</td>
<td align="center" valign="middle">4</td>
<td align="center" valign="middle">2&#x202F;mA</td>
<td align="left" valign="middle">1&#x202F;h<break/>online</td>
<td align="left" valign="middle">DLPFC<break/>anodes: F6/FC6<break/>cathodes: Fp2/AF8/AF4<break/>M1<break/>anodes: Cp1/Cp3<break/>cathodes: Fp1/F8/F9<break/>(15.7&#x202F;cm<sup>2</sup>)</td>
<td align="left" valign="middle">60&#x202F;s ramp up/60&#x202F;s ramp down</td>
<td align="left" valign="middle">Smoother landings during sessions 3 and 4 if active than sham over DLPFC only</td>
</tr>
<tr>
<td align="left" valign="middle">
<xref ref-type="bibr" rid="ref56">Mark et al. (2023)</xref>
</td>
<td align="left" valign="middle">Between groups</td>
<td align="left" valign="middle">Single-masked</td>
<td align="center" valign="middle"><italic>N</italic> =&#x202F;24<break/><italic>n</italic><sub>active</sub> <italic>=</italic> 12<break/>(<italic>n</italic><sub>novice active</sub> =&#x202F;6)<break/><italic>n</italic><sub>sham</sub> =&#x202F;12<break/>(<italic>n</italic><sub>novice sham</sub> =&#x202F;6)</td>
<td align="center" valign="middle">1</td>
<td align="center" valign="middle">1.5&#x202F;mA</td>
<td align="left" valign="middle">30&#x202F;min online</td>
<td align="left" valign="middle">Anode: AF8<break/>cathodes: Fpz/T8<break/>(8&#x202F;cm<sup>2</sup>)</td>
<td align="left" valign="middle">30&#x202F;s ramp up/30&#x202F;s ramp down</td>
<td align="left" valign="middle">Smoother landings if active than sham; stronger effects for novices than experts</td>
</tr>
<tr>
<td align="left" valign="middle">
<xref ref-type="bibr" rid="ref30">Feltman and Kelley (2024)</xref>
</td>
<td align="left" valign="middle">Mixed groups</td>
<td align="left" valign="middle">Single-masked</td>
<td align="center" valign="middle"><italic>N</italic> =&#x202F;22<break/><italic>n</italic><sub>online</sub> <italic>=</italic> 12<break/><italic>n</italic><sub>offline</sub> =&#x202F;10</td>
<td align="center" valign="middle">4</td>
<td align="center" valign="middle">2&#x202F;mA</td>
<td align="left" valign="middle">Online:2&#x00D7; 10&#x202F;min<break/>Offline:<break/>20&#x202F;min</td>
<td align="left" valign="middle">Anode: P4<break/>cathode: Fp1<break/>(25&#x202F;cm<sup>2</sup>)</td>
<td align="left" valign="middle">Online<break/>2&#x00D7; 30&#x202F;s ramp up/30&#x202F;s ramp down<break/>offline<break/>60&#x202F;s ramp up/60&#x202F;s ramp down</td>
<td align="left" valign="middle">More likely to follow glide path when active than sham for online only</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<p>DLPFC, dorsolateral prefrontal cortex; M1, motor cortex.</p>
</table-wrap-foot>
</table-wrap>
<sec id="sec3">
<label>2.1</label>
<title>tDCS and driving simulators</title>
<p>In the earliest study, <xref ref-type="bibr" rid="ref7">Beeli et al. (2008)</xref> examined the effects of tDCS over the DLPFC on driving metrics such as speed, headway distance, and lane positioning. Currently, outcomes measured in driving simulators lack gold standard metrics to define meaningful performance changes that translate to real-world driving. Recently, however, research has proposed two composite factors of driving behavior&#x2014;vehicle control variability and speed&#x2014;that include metrics like lane positioning, which also have strong face validity as indicators of safe driving (<xref ref-type="bibr" rid="ref58">McManus et al., 2024</xref>).</p>
<p>Across three sessions, <xref ref-type="bibr" rid="ref7">Beeli et al. (2008)</xref> tasked 21 participants (20&#x2013;30&#x202F;years, all men) to complete a 3-kilometer drive through a city scene with simulated traffic, lights and signs, and pedestrians. The first session was a baseline drive without tDCS. During the other two sessions, the anode and cathode were positioned unilaterally over the DLPFC in a counterbalanced order. Half of the participants randomly received stimulation over the left DLPFC at scalp coordinate F3, while the other half received stimulation over the right DLPFC at scalp coordinate F4. Stimulation was delivered at 1&#x202F;mA for 15&#x202F;min offline. Compared to the baseline, participants exhibited fewer speeding violations and maintained greater headway distance when receiving anodal tDCS over the DLPFC compared to cathodal. There were no observable effects of the hemisphere.</p>
<p>Several methodological considerations in this early study must be addressed. Without a sham condition or adequate blinding, it is difficult to disentangle stimulation effects from experimenter influence and participant bias (<xref ref-type="bibr" rid="ref15">Boutron et al., 2007</xref>). The most common sham procedure ramps up and down the current at the beginning and end of the protocol to mimic initial cutaneous sensations without lasting effects (<xref ref-type="bibr" rid="ref96">Woods et al., 2016</xref>). Additionally, a between-groups design introduces random variability (<xref ref-type="bibr" rid="ref14">Borghini et al., 2014</xref>; <xref ref-type="bibr" rid="ref49">Lakens, 2013</xref>) and fails to account for individual differences in tDCS effects, which can be influenced by anatomical factors (e.g., skull thickness) and behavioral baselines (<xref ref-type="bibr" rid="ref13">Bikson et al., 2012</xref>; <xref ref-type="bibr" rid="ref25">Datta et al., 2012</xref>; <xref ref-type="bibr" rid="ref37">Horvath et al., 2014</xref>; <xref ref-type="bibr" rid="ref44">Kim et al., 2014</xref>; <xref ref-type="bibr" rid="ref51">Li et al., 2015</xref>; <xref ref-type="bibr" rid="ref70">Opitz et al., 2015</xref>; <xref ref-type="bibr" rid="ref85">Splittgerber et al., 2020</xref>).</p>
<p><xref ref-type="bibr" rid="ref82">Sakai et al. (2014)</xref> conducted a sham-controlled, within-groups, single-masked study to address these limitations. Thirteen participants (~35&#x202F;years, 11 men) were instructed to maintain a specific headway distance from a lead vehicle over a 22-kilometer route. Participants completed this driving task over three testing sessions. In a counterbalanced order, participants received anodal tDCS over the right DLPFC at F4, cathodal tDCS at F4, and sham. Stimulation was set at 1.5&#x202F;mA for up to 20&#x202F;min online. There was less variability in headway distance and lane positioning when anodal tDCS was delivered over the DLPFC compared to cathodal and sham. This finding is consistent with research in other domains showing anodal-excitatory effects, but not cathodal-inhibitory effects, for cognitive tasks involving the DLPFC (<xref ref-type="bibr" rid="ref41">Jacobson et al., 2012</xref>). One explanation could be that the anode likely enhances neuronal firing in active areas, while the cathode may not sufficiently inhibit firing in highly active states.</p>
<p>The neuromodulation field has significantly advanced in the decade since <xref ref-type="bibr" rid="ref82">Sakai et al. (2014)</xref> published their work. <xref ref-type="bibr" rid="ref29">Facchin et al. (2023)</xref> explored the effects of different tDCS electrode montages on driving behavior in the latest driving study. Twenty-seven participants (21&#x2013;30&#x202F;years, 14 women) completed three 25-min driving sessions while receiving sham tDCS, conventional tDCS, or 4&#x202F;&#x00D7;&#x202F;1 HD-tDCS, where four electrodes surround a center electrode of the opposite polarity (<xref ref-type="bibr" rid="ref24">Datta et al., 2009</xref>; <xref ref-type="bibr" rid="ref48">Kuo et al., 2013</xref>). Anodal tDCS was applied over the right frontal eye field (FEF) at 1.5&#x202F;mA over FC4, an area implicated in visuomotor control (<xref ref-type="bibr" rid="ref19">Cameron et al., 2015</xref>; <xref ref-type="bibr" rid="ref35">Grosbras et al., 2005</xref>; <xref ref-type="bibr" rid="ref68">Nobre et al., 2000</xref>). Given that electrode size and material affect spatial resolution, coupled with the structural-functional connectivity of the human brain (<xref ref-type="bibr" rid="ref72">Park and Friston, 2013</xref>; <xref ref-type="bibr" rid="ref86">Sporns, 2013</xref>), the DLPFC may have been incidentally targeted during stimulation.</p>
<p>As many can attest, drivers rarely focus on just car following. To this point, <xref ref-type="bibr" rid="ref29">Facchin et al. (2023)</xref> manipulated driving task difficulty using two variations of stimulus&#x2013;response detection tasks commonly used in human factors research (<xref ref-type="bibr" rid="ref40">Innes et al., 2021</xref>). During the drive, the lead car frequently flashed its brake lights, and road signs appeared at random intervals. Participants were asked to brake in response to the lead vehicle and respond to the road signs. Outcomes measured included lane-keeping position, braking reaction time and accuracy, and road sign reaction time and accuracy. Lane maintenance was unaffected by stimulation. <xref ref-type="bibr" rid="ref29">Facchin et al. (2023)</xref> found that participants responded more quickly, though not more accurately, to the brake lights and road signs when receiving anodal tDCS over the FEF than sham. More prominent effects for these reaction times emerged when stimulation was delivered with HD-tDCS rather than conventional, suggesting heightened response speed to relevant stimuli. Together, these three driving studies indicate that anodal tDCS over the DLPFC may influence distance perception or judgment, observable as changes in distance or faster response times.</p>
</sec>
<sec id="sec4">
<label>2.2</label>
<title>tDCS and flight simulators</title>
<p><xref ref-type="bibr" rid="ref21">Choe et al. (2016)</xref> examined the impact of tDCS on skill acquisition and performance across various simulated flight tasks, using scenarios with computer-based simulations that align with Federal Aviation Administration (FAA) Industry Training Standards (FITS) to enhance real-world training relevance (<xref ref-type="bibr" rid="ref95">Williams, 2012</xref>). Though the performance was tested on flight tasks of varying difficulty, results were only published for the easiest task. Across four sessions, 32 participants (ages 21&#x2013;64, 31 men) attempted to replicate a landing demonstrated in an instructional video under daylight conditions with complete visibility. Measured outcomes included landing gravitational force (<italic>g</italic>-force), deviations from flight path, vertical speed, and vertical speed variance. While <italic>g</italic>-force assessment captures landing skill at the most challenging and critical phase of flight, the entire approach is considered with path and vertical speed deviations. Learning rates were measured across sessions, within sessions, and between trials or scenarios (5 trials per session).</p>
<p><xref ref-type="bibr" rid="ref21">Choe et al. (2016)</xref> treated stimulation application and location as between-group factors. Half the participants received anodal tDCS (2&#x202F;mA, 1&#x202F;h online), while the other half received sham. Stimulation was delivered over the right DLPFC (anodes F6/FC6) or the left motor cortex (anodes CP1/CP3). No significant effects emerged for the motor cortex group. In the DLPFC group, there was less variability in landing <italic>g</italic>-force observed during the third and fourth sessions, suggesting that tDCS may be more beneficial for trained tasks over time. <xref ref-type="bibr" rid="ref21">Choe et al. (2016)</xref> also collected EEG and functional near-infrared spectroscopy (fNIRS) data that suggests that participants who received active tDCS exhibited altered neuronal activity in the DLPFC and motor cortex compared to those who received sham. Interestingly, behavioral outcomes are commonly observed when delivering anodal excitation over the motor cortex, but not cognitive regions like the DLPFC (<xref ref-type="bibr" rid="ref41">Jacobson et al., 2012</xref>; <xref ref-type="bibr" rid="ref89">Tremblay et al., 2014</xref>). The broad influence of the DLPFC on cognitive functions, especially when considering varying stimulation parameters, makes predicting specific behavioral outcomes difficult.</p>
<p>This initial flight study was conducted under relatively simple conditions to facilitate task performance. However, more realistic scenarios may include bad weather, a narrow runway, or auditory distractions. Accordingly, <xref ref-type="bibr" rid="ref56">Mark et al. (2023)</xref> adjusted the workload during landing. Twenty-four glider pilots (ages 18&#x2013;22, mostly men) were recruited and categorized as novices or experts based on experience. Participants completed three runs in a single session, with a pre- and post-training run flanking a tDCS run. In the training run, participants received either sham or anodal tDCS over the right DLPFC at AF8 (1.5&#x202F;mA, 30&#x202F;min online). Feedback about performance was presented after each trial (72 trials in total). Measures included landing <italic>g</italic>-force, landing descent speed, and flair.</p>
<p><xref ref-type="bibr" rid="ref56">Mark et al. (2023)</xref> observed significant stimulation effects only for landing <italic>g</italic>-force. Specifically, participants who received active tDCS compared to those who received sham landed more smoothly when comparing pre-training to training and post-training. This skill-learning effect was more pronounced in novices than experts, similar to findings in electronic sports (<xref ref-type="bibr" rid="ref88">Toth et al., 2021</xref>), suggesting novices may benefit more from tDCS. The study took place in a functional magnetic resonance imaging (fMRI) machine, revealing that active tDCS than sham increases DLPFC activity and enhances connectivity between the DLPFC and cerebellum, a region involved in error-feedback learning (<xref ref-type="bibr" rid="ref27">Doya, 1999</xref>).</p>
<p>Targeting other brain regions with tDCS, such as the posterior parietal cortex (PPC), which guides the visuospatial orienting of selective attention (<xref ref-type="bibr" rid="ref8">Behrmann et al., 2004</xref>; <xref ref-type="bibr" rid="ref23">Culham and Valyear, 2006</xref>; <xref ref-type="bibr" rid="ref46">Kravitz et al., 2011</xref>; <xref ref-type="bibr" rid="ref53">Lo et al., 2019</xref>; <xref ref-type="bibr" rid="ref80">Rojas et al., 2018</xref>), could clarify the brain-behavior relationship in flight skill acquisition. In the most recent study, <xref ref-type="bibr" rid="ref30">Feltman and Kelley (2024)</xref> recruited 22 pilots (~37&#x202F;years, all men) to complete a 90-min round-trip flight while receiving anodal tDCS over the PPC at P4 (2&#x202F;mA, 20&#x202F;min total) and sham. Stimulation timing was treated as a between-groups condition between groups. Participants in the offline stimulation group received anodal tDCS (2&#x202F;mA, 20&#x202F;min) and sham before flight. Those in the online group received sham and anodal tDCS (2&#x202F;mA) delivered for 10&#x202F;min at 30 and 60&#x202F;min into the flight.</p>
<p>Toward the end of each flight leg, an emergency required participants to disengage autopilot and land safely. Altitude, airspeed, and heading were measured throughout the flight, while glideslope (vertical) and localizer (lateral) deviations were recorded during the approach. Significant effects emerged only for glideslope deviations in the online group, with online anodal tDCS associated with better alignment to the glide path than sham. These findings align with the role of the PPC in visuospatial attention. Together, these flight studies suggest that tDCS over the DLPFC and PPC may enhance landing smoothness, each investigating different aspects of stimulation and simulation parameters.</p>
</sec>
</sec>
<sec sec-type="discussion" id="sec5">
<label>3</label>
<title>Discussion</title>
<p>Operating a vehicle requires substantial cognitive, perceptual, and motor resources. Non-invasive neuromodulation methods, like tDCS, may offer insights into human performance when cognitive and perceptual enhancement are beneficial. This review synthesizes research on how targeting various brain regions via tDCS can influence outcomes in interactive driving and flight simulators.</p>
<p>Driving studies indicate that anodal tDCS over the DLPFC affects lateral and vertical lane positioning when following a lead vehicle (<xref ref-type="bibr" rid="ref7">Beeli et al., 2008</xref>; <xref ref-type="bibr" rid="ref29">Facchin et al., 2023</xref>; <xref ref-type="bibr" rid="ref82">Sakai et al., 2014</xref>). These findings suggest that tDCS can acutely impact operational (automatic, reactive) and maneuvering (controlled, tactical) driving behaviors (<xref ref-type="bibr" rid="ref60">Michon, 1985</xref>). It is also likely that tDCS can influence strategic (goal-directed, proactive) driving behaviors, such as trip planning, route memory, or adapting to detours (<xref ref-type="bibr" rid="ref60">Michon, 1985</xref>). Expanding the complexity of tasks to include strategic and goal-directed elements could be one approach to enhance functional and psychological fidelity, thereby bolstering task realism and immersion (<xref ref-type="bibr" rid="ref79">Roberts A. P. J. et al., 2020</xref>). Defining meaningful performance benchmarks in driving simulators can further aid in translating research findings into practical, everyday use (<xref ref-type="bibr" rid="ref58">McManus et al., 2024</xref>).</p>
<p>Similarly, the flight studies demonstrate that anodal tDCS over the DLPFC and PPC is associated with smoother landings, supported by converging neurophysiological evidence from EEG, fNIRS, and fMRI (<xref ref-type="bibr" rid="ref21">Choe et al., 2016</xref>; <xref ref-type="bibr" rid="ref30">Feltman and Kelley, 2024</xref>; <xref ref-type="bibr" rid="ref56">Mark et al., 2023</xref>). Although landing is one of the most challenging tasks for pilots, most of the time spent in flight involves monitoring system controls, including autopilot. For example, future studies may wish to explore the effects of tDCS during monotonous monitoring tasks. This inquiry becomes even more interesting when considering that visual scanning strategies are modulated by expertise (<xref ref-type="bibr" rid="ref50">Lefrancois et al., 2016</xref>; <xref ref-type="bibr" rid="ref54">Lounis et al., 2021</xref>). Combining stimulation with multimodal training may enhance its effects (<xref ref-type="bibr" rid="ref92">Ward et al., 2017</xref>) and contribute further to research on the long-term impacts of tDCS.</p>
<p>Several factors should be carefully considered when interpreting these findings and designing future research. Stimulation protocols must be optimized to reduce individual variability and potential biases. Within-group designs, sham controls, double-masking procedures, and carefully worded materials are some ways to address participant and experimenter biases. It is also critical to address systematic racial bias in neurophysiological research. Most studies in this review recruited small samples of young men, and participants&#x2019; race or ethnicity was not reported. This omission raises concerns about inclusivity and generalizability, as methods that require adherence between electrodes and the scalp often exclude individuals based on hair type and style (<xref ref-type="bibr" rid="ref22">Choy et al., 2022</xref>; <xref ref-type="bibr" rid="ref73">Parker and Ricard, 2022</xref>; <xref ref-type="bibr" rid="ref78">Roberts S. O. et al., 2020</xref>). Diverse, representative samples are essential to extend research beyond the lab and achieve broader inclusivity.</p>
<p>In summary, tDCS has the potential to modulate brain activity in regions that facilitate vehicle operation on the ground and in the clouds. To deepen our understanding of neuromodulation for human enhancement and continue exploring its possibilities, it is crucial to design stimulation protocols that mitigate biases and conduct studies with tasks or environments that reflect real-world conditions. As the promise of tDCS grows, it is essential to conduct rigorous investigations to fully understand its implications and optimize its application in various contexts.</p>
</sec>
</body>
<back>
<sec sec-type="author-contributions" id="sec6">
<title>Author contributions</title>
<p>KS: Conceptualization, Investigation, Methodology, Project administration, Validation, Visualization, Writing &#x2013; original draft, Writing &#x2013; review &#x0026; editing. NW: Conceptualization, Project administration, Supervision, Writing &#x2013; review &#x0026; editing.</p>
</sec>
<sec sec-type="funding-information" id="sec7">
<title>Funding</title>
<p>The author(s) declare that no financial support was received for the research, authorship, and/or publication of this article.</p>
</sec>
<sec sec-type="COI-statement" id="sec8">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec sec-type="disclaimer" id="sec9">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
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