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<front>
<journal-meta>
<journal-id journal-id-type="nlm-ta">PLoS ONE</journal-id>
<journal-id journal-id-type="publisher-id">plos</journal-id>
<journal-id journal-id-type="pmc">plosone</journal-id>
<journal-title-group>
<journal-title>PLOS ONE</journal-title>
</journal-title-group>
<issn pub-type="epub">1932-6203</issn>
<publisher>
<publisher-name>Public Library of Science</publisher-name>
<publisher-loc>San Francisco, CA USA</publisher-loc>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.1371/journal.pone.0220205</article-id>
<article-id pub-id-type="publisher-id">PONE-D-19-09978</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Research Article</subject>
</subj-group>
<subj-group subj-group-type="Discipline-v3"><subject>Medicine and health sciences</subject><subj-group><subject>Public and occupational health</subject><subj-group><subject>Physical activity</subject><subj-group><subject>Physical fitness</subject><subj-group><subject>Exercise</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Medicine and health sciences</subject><subj-group><subject>Sports and exercise medicine</subject><subj-group><subject>Exercise</subject></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and life sciences</subject><subj-group><subject>Sports science</subject><subj-group><subject>Sports and exercise medicine</subject><subj-group><subject>Exercise</subject></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and life sciences</subject><subj-group><subject>Neuroscience</subject><subj-group><subject>Cognitive science</subject><subj-group><subject>Cognitive psychology</subject><subj-group><subject>Creativity</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and life sciences</subject><subj-group><subject>Psychology</subject><subj-group><subject>Cognitive psychology</subject><subj-group><subject>Creativity</subject></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Social sciences</subject><subj-group><subject>Psychology</subject><subj-group><subject>Cognitive psychology</subject><subj-group><subject>Creativity</subject></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and life sciences</subject><subj-group><subject>Neuroscience</subject><subj-group><subject>Cognitive science</subject><subj-group><subject>Cognition</subject></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Research and analysis methods</subject><subj-group><subject>Mathematical and statistical techniques</subject><subj-group><subject>Statistical methods</subject><subj-group><subject>Regression analysis</subject></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Physical sciences</subject><subj-group><subject>Mathematics</subject><subj-group><subject>Statistics</subject><subj-group><subject>Statistical methods</subject><subj-group><subject>Regression analysis</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Research and analysis methods</subject><subj-group><subject>Bioassays and physiological analysis</subject><subj-group><subject>Electrophysiological techniques</subject><subj-group><subject>Cardiac electrophysiology</subject><subj-group><subject>Electrocardiography</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Medicine and health sciences</subject><subj-group><subject>Cardiology</subject><subj-group><subject>Heart rate</subject></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and life sciences</subject><subj-group><subject>Psychology</subject><subj-group><subject>Psychological adjustment</subject></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Social sciences</subject><subj-group><subject>Psychology</subject><subj-group><subject>Psychological adjustment</subject></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Medicine and health sciences</subject><subj-group><subject>Public and occupational health</subject><subj-group><subject>Physical activity</subject><subj-group><subject>Physical fitness</subject></subj-group></subj-group></subj-group></subj-group></article-categories>
<title-group>
<article-title>Creative challenge: Regular exercising moderates the association between task-related heart rate variability changes and individual differences in originality</article-title>
<alt-title alt-title-type="running-head">Task-related HRV changes and creative thinking</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes" xlink:type="simple">
<contrib-id authenticated="true" contrib-id-type="orcid">http://orcid.org/0000-0003-3195-4555</contrib-id>
<name name-style="western">
<surname>Rominger</surname>
<given-names>Christian</given-names>
</name>
<role content-type="http://credit.casrai.org/">Conceptualization</role>
<role content-type="http://credit.casrai.org/">Data curation</role>
<role content-type="http://credit.casrai.org/">Formal analysis</role>
<role content-type="http://credit.casrai.org/">Methodology</role>
<role content-type="http://credit.casrai.org/">Writing – original draft</role>
<xref ref-type="aff" rid="aff001"><sup>1</sup></xref>
<xref ref-type="corresp" rid="cor001">*</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Papousek</surname>
<given-names>Ilona</given-names>
</name>
<role content-type="http://credit.casrai.org/">Conceptualization</role>
<role content-type="http://credit.casrai.org/">Funding acquisition</role>
<role content-type="http://credit.casrai.org/">Methodology</role>
<role content-type="http://credit.casrai.org/">Project administration</role>
<role content-type="http://credit.casrai.org/">Writing – original draft</role>
<xref ref-type="aff" rid="aff001"><sup>1</sup></xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Fink</surname>
<given-names>Andreas</given-names>
</name>
<role content-type="http://credit.casrai.org/">Methodology</role>
<role content-type="http://credit.casrai.org/">Writing – review &amp; editing</role>
<xref ref-type="aff" rid="aff001"><sup>1</sup></xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Perchtold</surname>
<given-names>Corinna M.</given-names>
</name>
<role content-type="http://credit.casrai.org/">Data curation</role>
<role content-type="http://credit.casrai.org/">Methodology</role>
<role content-type="http://credit.casrai.org/">Writing – review &amp; editing</role>
<xref ref-type="aff" rid="aff001"><sup>1</sup></xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple">
<contrib-id authenticated="true" contrib-id-type="orcid">http://orcid.org/0000-0002-0159-3720</contrib-id>
<name name-style="western">
<surname>Lackner</surname>
<given-names>Helmut K.</given-names>
</name>
<role content-type="http://credit.casrai.org/">Methodology</role>
<role content-type="http://credit.casrai.org/">Writing – review &amp; editing</role>
<xref ref-type="aff" rid="aff002"><sup>2</sup></xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Weiss</surname>
<given-names>Elisabeth M.</given-names>
</name>
<role content-type="http://credit.casrai.org/">Methodology</role>
<role content-type="http://credit.casrai.org/">Writing – review &amp; editing</role>
<xref ref-type="aff" rid="aff001"><sup>1</sup></xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple">
<contrib-id authenticated="true" contrib-id-type="orcid">http://orcid.org/0000-0002-0371-3730</contrib-id>
<name name-style="western">
<surname>Schwerdtfeger</surname>
<given-names>Andreas R.</given-names>
</name>
<role content-type="http://credit.casrai.org/">Conceptualization</role>
<role content-type="http://credit.casrai.org/">Methodology</role>
<role content-type="http://credit.casrai.org/">Writing – original draft</role>
<xref ref-type="aff" rid="aff001"><sup>1</sup></xref>
</contrib>
</contrib-group>
<aff id="aff001"><label>1</label> <addr-line>Department of Psychology, University of Graz, Graz, Austria</addr-line></aff>
<aff id="aff002"><label>2</label> <addr-line>Otto Loewi Research Center, Section of Physiology, Medical University of Graz, Graz, Austria</addr-line></aff>
<contrib-group>
<contrib contrib-type="editor" xlink:type="simple">
<name name-style="western">
<surname>Boullosa</surname>
<given-names>Daniel</given-names>
</name>
<role>Editor</role>
<xref ref-type="aff" rid="edit1"/>
</contrib>
</contrib-group>
<aff id="edit1"><addr-line>University of Brasilia, BRAZIL</addr-line></aff>
<author-notes>
<fn fn-type="conflict" id="coi001">
<p>The authors have declared that no competing interests exist.</p>
</fn>
<corresp id="cor001">* E-mail: <email xlink:type="simple">christian.rominger@uni-graz.at</email></corresp>
</author-notes>
<pub-date pub-type="epub">
<day>22</day>
<month>7</month>
<year>2019</year>
</pub-date>
<pub-date pub-type="collection">
<year>2019</year>
</pub-date>
<volume>14</volume>
<issue>7</issue>
<elocation-id>e0220205</elocation-id>
<history>
<date date-type="received">
<day>8</day>
<month>4</month>
<year>2019</year>
</date>
<date date-type="accepted">
<day>10</day>
<month>7</month>
<year>2019</year>
</date>
</history>
<permissions>
<copyright-year>2019</copyright-year>
<copyright-holder>Rominger et al</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">
<license-p>This is an open access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">Creative Commons Attribution License</ext-link>, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
</license>
</permissions>
<self-uri content-type="pdf" xlink:href="info:doi/10.1371/journal.pone.0220205"/>
<abstract>
<p>Coping with mental challenges is vital to everyday functioning. In accordance with prominent theories, the adaptive and flexible adjustment of the organism to daily demands is well expressed in task-related changes of cardiac vagal control. While many mental challenges are associated with increased effort and associated decreased task-related heart rate variability (HRV), some cognitive challenges go along with HRV increases. Especially creativity represents a cognitive process, which not only results from mental effort but also from spontaneous modes of thinking. Critically, creativity and HRV are associated with regular exercising and fitness. Furthermore, the cross-stressor adaptation theory suggests that changes in cardiac reactions to physical challenges may generalize to mental challenges. In line with this idea the amount of regular exercising was hypothesized to moderate the association between HRV changes and creativity. A sample of 97 participants was investigated. They reported the amount of regular exercise and their ECG was measured at baseline and during a creativity task. An association between task-related HRV changes and originality as a function of participants’ amount of regular exercise was found. Participants reporting more regular exercising produced more original ideas when they had higher HRV increases during the task, while more sedentary participants showed the opposite association. Results suggest that individuals with a higher amount of regular exercise achieve higher originality probably via the engagement in more spontaneous modes of thinking, while more sedentary people may primarily benefit from increased mental effort. This supports the conclusion that higher creativity can be achieved by different strategies.</p>
</abstract>
<funding-group>
<award-group id="award001">
<funding-source>
<institution-wrap>
<institution-id institution-id-type="funder-id">http://dx.doi.org/10.13039/501100002428</institution-id>
<institution>Austrian Science Fund</institution>
</institution-wrap>
</funding-source>
<award-id>P 30362</award-id>
<principal-award-recipient>
<name name-style="western">
<surname>Papousek</surname>
<given-names>Ilona</given-names>
</name>
</principal-award-recipient>
</award-group>
<funding-statement>This research was supported by a grant from the Austrian Science Fund (FWF): P 30362, awarded to I.P. (<ext-link ext-link-type="uri" xlink:href="https://pf.fwf.ac.at/de/wissenschaft-konkret/project-finder/?search%5Bwhat%5D=P+30362&amp;search%5Bscience_discipline_id%5D=&amp;search%5Bpromotion_category_id%5D=&amp;extended=1" xlink:type="simple">https://pf.fwf.ac.at/de/wissenschaft-konkret/project-finder/?search%5Bwhat%5D=P+30362&amp;search%5Bscience_discipline_id%5D=&amp;search%5Bpromotion_category_id%5D=&amp;extended=1</ext-link>). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.</funding-statement>
</funding-group>
<counts>
<fig-count count="3"/>
<table-count count="1"/>
<page-count count="14"/>
</counts>
<custom-meta-group>
<custom-meta id="data-availability">
<meta-name>Data Availability</meta-name>
<meta-value>All relevant data are within the paper and its Supporting Information files.</meta-value>
</custom-meta>
</custom-meta-group>
</article-meta>
</front>
<body>
<sec id="sec001" sec-type="intro">
<title>Introduction</title>
<p>Diverse mental challenges are a major part of daily life, and the flexible adjustment to these day-to-day demands seems mandatory for successful coping [<xref ref-type="bibr" rid="pone.0220205.ref001">1</xref>]. However, this adaptive ability of self-regulation greatly varies between people and situations. Notably, mental effort, stress, and self-regulation are indexed by functions of the vagus nerve as the primary part of the parasympathetic nervous system [<xref ref-type="bibr" rid="pone.0220205.ref001">1</xref>–<xref ref-type="bibr" rid="pone.0220205.ref005">5</xref>]. A putative physiological marker of vagal control is the beat to beat variation in heart rate, the so-called heart rate variability (HRV) [<xref ref-type="bibr" rid="pone.0220205.ref006">6</xref>,<xref ref-type="bibr" rid="pone.0220205.ref007">7</xref>]. The vagal influence on HRV can be quantified in the time domain by means of RMSSD (root mean square of the successive difference in normal sinus beat intervals) and pNN50 (percentage of adjacent normal sinus beat intervals that differ by more than 50 ms) [<xref ref-type="bibr" rid="pone.0220205.ref008">8</xref>–<xref ref-type="bibr" rid="pone.0220205.ref010">10</xref>].</p>
<p>While the tonic HRV level potentially indicates an inter-individual trait like capacity to adapt to environmental challenges by parasympathetic/vagal control [<xref ref-type="bibr" rid="pone.0220205.ref011">11</xref>–<xref ref-type="bibr" rid="pone.0220205.ref013">13</xref>] (for a meta-analysis see [<xref ref-type="bibr" rid="pone.0220205.ref014">14</xref>]), task-related HRV changes represent the dynamic task-specific vagal control of the heart (i.e., vagal break) [<xref ref-type="bibr" rid="pone.0220205.ref002">2</xref>,<xref ref-type="bibr" rid="pone.0220205.ref015">15</xref>]. Therefore, task-related HRV is an even more sensitive variable in the context of regulatory efforts of the organism and the actual flexible adaption to physical and mental challenges [<xref ref-type="bibr" rid="pone.0220205.ref002">2</xref>,<xref ref-type="bibr" rid="pone.0220205.ref003">3</xref>,<xref ref-type="bibr" rid="pone.0220205.ref006">6</xref>,<xref ref-type="bibr" rid="pone.0220205.ref007">7</xref>,<xref ref-type="bibr" rid="pone.0220205.ref013">13</xref>,<xref ref-type="bibr" rid="pone.0220205.ref016">16</xref>,<xref ref-type="bibr" rid="pone.0220205.ref017">17</xref>]. Following from this, the adjustment of the organism to task-related challenges—e.g., self-regulatory efforts [<xref ref-type="bibr" rid="pone.0220205.ref004">4</xref>,<xref ref-type="bibr" rid="pone.0220205.ref013">13</xref>]—may be best represented by the flexible change of the HRV from a baseline to a period of mental effort.</p>
<p>However, not every mental challenge requires the same amount of cognitive and regulatory capacity [<xref ref-type="bibr" rid="pone.0220205.ref018">18</xref>,<xref ref-type="bibr" rid="pone.0220205.ref019">19</xref>]. Of note, while many mental tasks are associated with task-related HRV decreases [<xref ref-type="bibr" rid="pone.0220205.ref016">16</xref>,<xref ref-type="bibr" rid="pone.0220205.ref020">20</xref>,<xref ref-type="bibr" rid="pone.0220205.ref021">21</xref>], there is evidence that some cognitive tasks are associated with task-related increases [<xref ref-type="bibr" rid="pone.0220205.ref003">3</xref>,<xref ref-type="bibr" rid="pone.0220205.ref004">4</xref>,<xref ref-type="bibr" rid="pone.0220205.ref015">15</xref>]. For instance, Butler et al. [<xref ref-type="bibr" rid="pone.0220205.ref013">13</xref>] found HRV increases during the reappraisal of negative events [<xref ref-type="bibr" rid="pone.0220205.ref022">22</xref>,<xref ref-type="bibr" rid="pone.0220205.ref023">23</xref>], and Silvia et al. [<xref ref-type="bibr" rid="pone.0220205.ref024">24</xref>] reported a slight rise of vagal control during a creative ideation task. This is remarkable, since creative ideation and cognitive reappraisal (the latter as an instance of “creativity in an affective context” [<xref ref-type="bibr" rid="pone.0220205.ref025">25</xref>]) represent complex self-regulatory cognitive processes (e.g., flexible switch between associative and executive functions), which share important basic cognitive functions [<xref ref-type="bibr" rid="pone.0220205.ref025">25</xref>–<xref ref-type="bibr" rid="pone.0220205.ref028">28</xref>].</p>
<p>A unique characteristic of creative and divergent thinking tasks in contrast to traditional convergent cognitive tasks is the interplay of associative/spontaneous modes of thinking and executive top-down control processes [<xref ref-type="bibr" rid="pone.0220205.ref029">29</xref>–<xref ref-type="bibr" rid="pone.0220205.ref034">34</xref>]. Following dual process theories, creative ideation is not solely based on mental effort, which is associated with HRV decrease [<xref ref-type="bibr" rid="pone.0220205.ref024">24</xref>,<xref ref-type="bibr" rid="pone.0220205.ref035">35</xref>–<xref ref-type="bibr" rid="pone.0220205.ref038">38</xref>], but also on spontaneous, automatic, associative, and relaxed modes of thinking, which are presumably associated with HRV increase [<xref ref-type="bibr" rid="pone.0220205.ref039">39</xref>–<xref ref-type="bibr" rid="pone.0220205.ref042">42</xref>]. Furthermore, the flexible shift between these different modes of thinking is important for creative ideation performance [<xref ref-type="bibr" rid="pone.0220205.ref043">43</xref>–<xref ref-type="bibr" rid="pone.0220205.ref045">45</xref>]. The contribution of both modes of thinking to creative ideation may explain contradictory findings in literature. Loudon and Deininger [<xref ref-type="bibr" rid="pone.0220205.ref035">35</xref>] reported a negative and Bowers and Keeling [<xref ref-type="bibr" rid="pone.0220205.ref040">40</xref>] a positive association between indices of creative performance and HRV. Silvia et al. [<xref ref-type="bibr" rid="pone.0220205.ref024">24</xref>] found no significant link between task-related vagal control and originality.</p>
<p>However, a further reason for inconsistent findings in this context might be that cognitive performance and HRV are associated with the amount of regular physical exercise, physical fitness, and general health of people [<xref ref-type="bibr" rid="pone.0220205.ref006">6</xref>,<xref ref-type="bibr" rid="pone.0220205.ref008">8</xref>,<xref ref-type="bibr" rid="pone.0220205.ref046">46</xref>] (for cognition see [<xref ref-type="bibr" rid="pone.0220205.ref047">47</xref>,<xref ref-type="bibr" rid="pone.0220205.ref048">48</xref>]; but see [<xref ref-type="bibr" rid="pone.0220205.ref049">49</xref>]; for cognition and HRV see [<xref ref-type="bibr" rid="pone.0220205.ref050">50</xref>]). Furthermore, in a recent study, Latorre Román et al. [<xref ref-type="bibr" rid="pone.0220205.ref051">51</xref>] found an association between measures of physical fitness and creativity (see also [<xref ref-type="bibr" rid="pone.0220205.ref052">52</xref>,<xref ref-type="bibr" rid="pone.0220205.ref053">53</xref>]; similar findings for acute physical activity see, [<xref ref-type="bibr" rid="pone.0220205.ref054">54</xref>–<xref ref-type="bibr" rid="pone.0220205.ref056">56</xref>]). The cross-stressor adaptation theory [<xref ref-type="bibr" rid="pone.0220205.ref057">57</xref>] assumes that salutary cardiovascular reactivity and recovery due to physical training generalize from ergogenic challenges to psychogenic challenges. This generalization may lead to a differentiation of cardiac reactivity to psychological stressors between people doing more and people doing less exercise and people who might therefore be more or less physically fit [<xref ref-type="bibr" rid="pone.0220205.ref046">46</xref>,<xref ref-type="bibr" rid="pone.0220205.ref058">58</xref>–<xref ref-type="bibr" rid="pone.0220205.ref060">60</xref>].</p>
<p>Following this idea, the present study examined whether regular exercising may moderate the association between task-related HRV changes and the performance outcome in a creative ideation task. People regularly exercising more are supposed to be physically best prepared to show adaptive task-related changes of HRV [<xref ref-type="bibr" rid="pone.0220205.ref006">6</xref>], in order to flexibly match the adaptive states important for optimum performance outcome [<xref ref-type="bibr" rid="pone.0220205.ref001">1</xref>]. More precisely, it was assumed that participants with a higher amount of regular exercise may more flexibly adjust to the creative challenge and more easily switch between mental effort [<xref ref-type="bibr" rid="pone.0220205.ref024">24</xref>,<xref ref-type="bibr" rid="pone.0220205.ref035">35</xref>] and spontaneous and automatic modes of thinking [<xref ref-type="bibr" rid="pone.0220205.ref040">40</xref>–<xref ref-type="bibr" rid="pone.0220205.ref042">42</xref>]. Therefore, it was hypothesized that participants exercising more often will show higher task-related HRV during creative ideation in contrast to participants exercising less, and that in exercisers (but not necessarily in sedentary people) the HRV response will be related to task performance.</p>
</sec>
<sec id="sec002" sec-type="materials|methods">
<title>Methods</title>
<sec id="sec003">
<title>Participants</title>
<p>An a-priori power analysis was conducted using the software G*Power 3.1 [<xref ref-type="bibr" rid="pone.0220205.ref061">61</xref>]. The analysis indicated that a sample size of 99 participants is required to detect a medium effect (<italic>f</italic><sup><italic>2</italic></sup> = .15) with a power of .80 and an alpha error of 5%. One hundred and two people were recruited to participated in the study because of potential technical failures and dropout from the study. Due to recording problems of the electrocardiogram two participants were excluded. Two further participants did not comply with the instructions, and another participant was excluded because of excessive artifacts in the recorded ECG. The final sample consisted of 97 participants (54 women) with an age range from 18 to 33 years (<italic>M</italic> = 23.07 years; <italic>SD</italic> = 3.48 years). Participants’ age range was restricted to control for potentially confounding influences on HRV [<xref ref-type="bibr" rid="pone.0220205.ref015">15</xref>].</p>
<p>People with a history of major psychiatric disorders according to the Structured Clinical Interview for DSM-IV Axis I Disorders (SCID) and people, who reported having a neurological disease or using psychoactive medication were not included in the study. All participants were right-handed (assessed by a standardized hand skill test; [<xref ref-type="bibr" rid="pone.0220205.ref062">62</xref>]). They were requested to refrain from alcohol intake for 12 h and from drinking coffee and other stimulating beverages for 2 h prior to their appointment, and to come to the session well rested. The study was approved by the Ethics Committee of the University of Graz, approval number GZ. 39/41/63 ex 2015/16. Written informed consent was obtained from all participants.</p>
</sec>
<sec id="sec004">
<title>Amount of regular physical exercise</title>
<p>The Freiburger Questionnaire on Physical Activity (FQPA) [<xref ref-type="bibr" rid="pone.0220205.ref063">63</xref>] is a reliable and valid self-report instrument, which covers everyday physical activities (e.g., walking to work), leisure time activities (e.g., dancing), and sports activities (e.g., swimming) [<xref ref-type="bibr" rid="pone.0220205.ref064">64</xref>,<xref ref-type="bibr" rid="pone.0220205.ref065">65</xref>]. Participants are instructed to indicate the duration for each activity in minutes per week which are converted into metabolic equivalents (MET) [<xref ref-type="bibr" rid="pone.0220205.ref066">66</xref>]. In order to derive a measure of physical exercise potentially resulting in higher bodily fitness [<xref ref-type="bibr" rid="pone.0220205.ref067">67</xref>], only the reported hours of sports activity per week (e.g., running, playing soccer, swimming, <italic>M</italic> = 3.41 h, <italic>SD</italic> = 3.68 h) were considered (exercise MET: <italic>M</italic> = 22.16, <italic>SD</italic> = 26.12) [<xref ref-type="bibr" rid="pone.0220205.ref068">68</xref>]. Frey et al. [<xref ref-type="bibr" rid="pone.0220205.ref063">63</xref>] reported a re-test reliability of <italic>r</italic> = .98 (two weeks) for the amount of exercise per week.</p>
</sec>
<sec id="sec005">
<title>Creative thinking task and task performance</title>
<p>The Alternate Uses (AU) task [<xref ref-type="bibr" rid="pone.0220205.ref069">69</xref>] is a verbal creativity/divergent thinking test used in numerous scientific studies [<xref ref-type="bibr" rid="pone.0220205.ref024">24</xref>,<xref ref-type="bibr" rid="pone.0220205.ref030">30</xref>,<xref ref-type="bibr" rid="pone.0220205.ref070">70</xref>]. The participants’ task was to generate alternative uses of common objects (e.g., umbrella, brick, or key). For the present investigation, a single answer (i.e., best idea) and self-paced version of the AU task was used. This approach was applied in order to more strongly focus on the originality aspect of creativity [<xref ref-type="bibr" rid="pone.0220205.ref026">26</xref>]. The self-paced procedure appropriately captures the spontaneous nature of the creative thinking process [<xref ref-type="bibr" rid="pone.0220205.ref071">71</xref>–<xref ref-type="bibr" rid="pone.0220205.ref075">75</xref>]; however, it also implicates high demands on self-regulatory mechanisms. Each trial started with a white cross (10 s), followed by a picture of a common object (idea generation phase with a max. response time of 15 s; see Perchtold et al. [<xref ref-type="bibr" rid="pone.0220205.ref026">26</xref>] for a similar procedure). After the “idea button” was pressed, the participants rated the originality of their idea on a 6-point Likert-scale (max. response time of 4 s) and finally voiced the idea (10 sec). The subjective rating was included to maintain the participants’ effort to produce high quality ideas for every single item of the task. With the presentation of a new fixation cross the next trial started (see <xref ref-type="fig" rid="pone.0220205.g001">Fig 1</xref> for an illustrative summary of the time course of the task). Sixteen objects were presented in randomized order.</p>
<fig id="pone.0220205.g001" position="float">
<object-id pub-id-type="doi">10.1371/journal.pone.0220205.g001</object-id>
<label>Fig 1</label>
<caption>
<title/>
<p>Schematic time course of the computerized (A) baseline, (B) AU task, and (C) the analysed 2 minutes time periods (Baseline, AU-ACT1, AU-ACT2, AU-ACT3).</p>
</caption>
<graphic mimetype="image" position="float" xlink:href="info:doi/10.1371/journal.pone.0220205.g001" xlink:type="simple"/>
</fig>
<p>The voiced ideas were transcribed and rated by three independent and trained raters on a four-point Likert scale ranging from “not original” to “very original”. This procedure is a common approach in creativity research (cf. Consensual Assessment Technique; [<xref ref-type="bibr" rid="pone.0220205.ref030">30</xref>,<xref ref-type="bibr" rid="pone.0220205.ref076">76</xref>]). The originality ratings showed acceptable interrater reliability (ICC (2, k) = .66).</p>
</sec>
<sec id="sec006">
<title>Procedure</title>
<p>All participants were tested individually. The recording took place in a separate and quiet room, where participants were seated in a comfortable chair. During the recordings, they were monitored via a web cam to ensure that they were following the instructions. After filling in the consent form, participants filled in the questionnaire assessing the amount of regular physical exercise. Then, the ECG was applied, and the baseline was recorded for two minutes. Participants were told to relax and keep their eyes closed until they heard a signal tone. After that, participants worked on tasks not relevant for the study question, which lasted a minimum of 10 minutes (see <xref ref-type="fig" rid="pone.0220205.g001">Fig 1</xref>). Then, the AU task followed.</p>
</sec>
<sec id="sec007">
<title>Recording and quantification of HRV</title>
<p>The electrocardiogram was recorded using a standard limb lead II electrode configuration with a sampling rate of 500 Hz (Brainvision Research Amplifier, Brain Products<sup>TM</sup>). The ECG signals were manually checked for artifacts by means of the software Kubios HRV Premium version 3.0.2. The mean heart rate of the participants at the baseline level was 68.61 bpm (<italic>SD</italic> = 11.11 bpm). HRV was quantified in the time domain by means of the root mean square of successive differences of the successive RR intervals. RMSSD has been shown to sensitively index vagal efference [<xref ref-type="bibr" rid="pone.0220205.ref077">77</xref>] especially during cognitive processes [<xref ref-type="bibr" rid="pone.0220205.ref021">21</xref>]. RMSSD was calculated for the two minutes baseline interval and three consecutive two minutes periods during the creative ideation task, starting from the onset of the first white cross (see <xref ref-type="fig" rid="pone.0220205.g001">Fig 1</xref>). Two minutes are considered adequate to evaluate RMSSD reliably [<xref ref-type="bibr" rid="pone.0220205.ref009">9</xref>,<xref ref-type="bibr" rid="pone.0220205.ref015">15</xref>,<xref ref-type="bibr" rid="pone.0220205.ref078">78</xref>]. As depicted in <xref ref-type="fig" rid="pone.0220205.g001">Fig 1</xref>, the three two minutes periods of the creative thinking task were calculated for comparability reasons with baseline ([<xref ref-type="bibr" rid="pone.0220205.ref009">9</xref>,<xref ref-type="bibr" rid="pone.0220205.ref015">15</xref>]; similar procedure see, [<xref ref-type="bibr" rid="pone.0220205.ref024">24</xref>,<xref ref-type="bibr" rid="pone.0220205.ref060">60</xref>,<xref ref-type="bibr" rid="pone.0220205.ref079">79</xref>]). The pooled untransformed RMSSD value of the three activation periods was <italic>M</italic> = 49.43 ms (<italic>SD</italic> = 24.57 ms) and the baseline was <italic>M</italic> = 51.63 ms (<italic>SD</italic> = 27.25 ms). The RMSSD reactivity showed high variability, with both increases and decreases (<italic>M</italic> = -2.20 ms, <italic>SD</italic> = 13.24 ms, min = -49.13 ms, max = 28.12 ms).</p>
<p>Furthermore, to control for respiration-induced changes in HRV, ECG derived respiration (EDR) was calculated by means of software Kubios HRV Premium version 3.0.2 for the baseline (<italic>M</italic> = 0.23 Hz, <italic>SD</italic> = 0.04 Hz) and the three consecutive two minutes activation periods, which were pooled together (<italic>M</italic> = 0.23 Hz, <italic>SD</italic> = 0.03 Hz).</p>
<p>On average, participants completed 11.73 (<italic>SD</italic> = 1.78) items during the three activation periods of overall 6 minutes (min = 6 items, max = 14 items). The range of response times from item onset to activation of the “idea button” was between 2933 ms and 12 329 ms (<italic>M</italic> = 6522 ms, <italic>SD</italic> = 2231 ms).</p>
</sec>
<sec id="sec008">
<title>Statistical analysis</title>
<p>Because of the skewed distribution of the RMSSD values, which was visually checked, a transformation with the natural logarithm (ln) was performed for all conducted statistical analyses [<xref ref-type="bibr" rid="pone.0220205.ref015">15</xref>,<xref ref-type="bibr" rid="pone.0220205.ref080">80</xref>–<xref ref-type="bibr" rid="pone.0220205.ref082">82</xref>].</p>
<p>Firstly, to explore the overall challenging effect of the creative thinking task, a one-way analysis of variance with the within-subjects factor TIME (Baseline, AU-ACT1, AU-ACT2, AU-ACT3) and the HRV (lnRMSSD) as the dependent variable was calculated. Due to violations of sphericity assumptions, the multivariate approach was used [<xref ref-type="bibr" rid="pone.0220205.ref083">83</xref>]. To illustrate the interindividual differences in HRV changes, the untransformed RMSSD values were presented.</p>
<p>The main research question, if regular exercising moderates the association between task-related HRV changes and creative ideation performance outcome, was evaluated using standard multiple regression analysis with the task-related HRV change score, the amount of physical exercise, and the interaction term as predictors, and task performance (originality) as the dependent variable. The task-related HRV change score was calculated by regressing the baseline lnRMSSD values on the pooled lnRMSSD values of the three two minutes activation periods (AU-ACT1, AU-ACT2, AU-ACT3) [<xref ref-type="bibr" rid="pone.0220205.ref084">84</xref>]. This procedure results in one residualized change index with higher scores indicating a relative increase in HRV and lower scores indicating a relative decrease in HRV during the creative ideation task. To illustrate the significant interaction effect of physical exercise and task-related HRV changes on originality, predicted originality was calculated for one standard deviation below sample mean (M—1 SD) and one standard deviation above sample mean (M + 1 SD) using standard regression analysis (similar procedure see e.g., [<xref ref-type="bibr" rid="pone.0220205.ref085">85</xref>,<xref ref-type="bibr" rid="pone.0220205.ref086">86</xref>]). To control for respiratory-induced changes in HRV, this standard multiple regression analysis was re-run with the mean EDR frequency of the three activation periods as an additional predictor.</p>
<p>All analyses were calculated by means of IBM SPSS Statistics 25 for windows and the significance level was set to 5%.</p>
</sec>
</sec>
<sec id="sec009" sec-type="results">
<title>Results</title>
<sec id="sec010">
<title>Overall effect of the creative thinking task on HRV</title>
<p>The analysis slightly failed to reach a significant main effect for TIME (<italic>F</italic>(3,94) = 2.45, <italic>p</italic> = .069, <italic>η</italic><sub><italic>p</italic></sub><sup><italic>2</italic></sup> = .03). As indicated in <xref ref-type="fig" rid="pone.0220205.g002">Fig 2</xref>, the strongest decrease of HRV was from baseline to the first interval of the creative ideation task.</p>
<fig id="pone.0220205.g002" position="float">
<object-id pub-id-type="doi">10.1371/journal.pone.0220205.g002</object-id>
<label>Fig 2</label>
<caption>
<title>Mean HRV (lnRMSSD; error bars = <italic>SE</italic>) in the baseline and the three consecutive activation (ACT) intervals during the AU task (AU-ACT1, AU-ACT2, AU-ACT3).</title>
</caption>
<graphic mimetype="image" position="float" xlink:href="info:doi/10.1371/journal.pone.0220205.g002" xlink:type="simple"/>
</fig>
</sec>
<sec id="sec011">
<title>Association between task-related HRV (lnRMSSD) change and task performance (originality), and its moderation by the amount of regular exercise</title>
<p>The regression analysis was significant (<italic>F</italic>(3,93) = 4.50, <italic>p =</italic> .005, <italic>R</italic><sup><italic>2</italic></sup> = .127). As depicted in <xref ref-type="table" rid="pone.0220205.t001">Table 1</xref>, the interaction effect of task-related HRV change by amount of physical exercise was significant (<italic>β</italic> = .34, <italic>p</italic> = .001), however the main effects of the amount of exercise and HRV change were non-significant. Please note that the results of the multiple regression analysis remained virtually the same when using pNN50 instead of RMSSD (<italic>F</italic>(3,93) = 2.98, <italic>p =</italic> .035, <italic>R</italic><sup><italic>2</italic></sup> = .09).</p>
<table-wrap id="pone.0220205.t001" position="float">
<object-id pub-id-type="doi">10.1371/journal.pone.0220205.t001</object-id>
<label>Table 1</label> <caption><title>Summary of the results of the multiple regression analysis for the prediction of task performance (originality) with task-related HRV (lnRMSSD) change, physical exercise, and the interaction term as predictors.</title></caption>
<alternatives>
<graphic id="pone.0220205.t001g" mimetype="image" position="float" xlink:href="info:doi/10.1371/journal.pone.0220205.t001" xlink:type="simple"/>
<table>
<colgroup>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
</colgroup>
<thead>
<tr>
<th align="left"/>
<th align="left"><italic>β</italic></th>
<th align="left"><italic>r</italic></th>
</tr>
</thead>
<tbody>
<tr>
<td align="left">Task-related HRV (lnRMSSD) change</td>
<td align="left">.06 (.538)</td>
<td align="left">.02 (.834)</td>
</tr>
<tr>
<td align="left">Physical exercise</td>
<td align="left">-.09 (.381)</td>
<td align="left">-.13 (.194)</td>
</tr>
<tr>
<td align="left">Task-related HRV (lnRMSSD) change x Physical exercise</td>
<td align="left">.34 (.001)</td>
<td align="left">.34 (.001)</td>
</tr>
</tbody>
</table>
</alternatives>
<table-wrap-foot>
<fn id="t001fn001"><p><italic>Note</italic>. <italic>p</italic>-values in parenthesis. <italic>r</italic> represents the zero-order correlation and <italic>β</italic> the standardized Beta coefficient.</p></fn>
</table-wrap-foot>
</table-wrap>
<p>To further qualify the interaction effect in the whole sample, the regression analysis was re-run with the amount of regular exercise set as low (M—1 SD) and high (M + 1 SD; simple slopes approach), respectively. As illustrated in <xref ref-type="fig" rid="pone.0220205.g003">Fig 3</xref>, participants exercising more often performed better in the creative thinking task in terms of originality when they showed greater relative HRV increases during the challenge (<italic>β</italic> = .40, <italic>t</italic>(93) = -2.73, <italic>p</italic> = .008). In participants exercising less this association was reversed (<italic>β</italic> = -.28, <italic>t</italic>(93) = -2.13, <italic>p</italic> = .036).</p>
<fig id="pone.0220205.g003" position="float">
<object-id pub-id-type="doi">10.1371/journal.pone.0220205.g003</object-id>
<label>Fig 3</label>
<caption>
<title>Interaction effect of task-related HRV change and the amount of regular physical exercise on task performance in terms of originality.</title>
</caption>
<graphic mimetype="image" position="float" xlink:href="info:doi/10.1371/journal.pone.0220205.g003" xlink:type="simple"/>
</fig>
<p>The regression analysis with the additional predictor EDR frequency was again significant (<italic>F</italic>(4,92) = 4.20, <italic>p =</italic> .004, <italic>R</italic><sup><italic>2</italic></sup> = .155). The interaction effect of task-related HRV change by amount of physical exercise remained significant (<italic>β</italic> = .32, <italic>p</italic> = .001) and the EDR frequency was not significant (<italic>β</italic> = .13, <italic>p</italic> = .214).</p>
<p>The correlation between the originality score and the number of generated ideas during the six minutes activation period was <italic>r</italic> = .05 (<italic>p</italic> = .596), indicating that the effects of the regression analysis was also not influenced by the number of generated ideas.</p>
</sec>
</sec>
<sec id="sec012" sec-type="conclusions">
<title>Discussion</title>
<p>The aim of this study was to examine the interplay of regular physical exercise and HRV reactivity during a challenging creative ideation task on task performance. Generally, HRV tended to decrease during the task, particularly at the beginning, which indicates the mobilization of mental effort and is characteristic of challenging performance tasks [<xref ref-type="bibr" rid="pone.0220205.ref087">87</xref>,<xref ref-type="bibr" rid="pone.0220205.ref088">88</xref>]. However, participants greatly varied in their cardiac response to the challenge. While some showed marked decreases of HRV, others exhibited smaller decreases or even increases of HRV during the task [<xref ref-type="bibr" rid="pone.0220205.ref015">15</xref>]. This variability in the cardiac responses was meaningfully related to task performance in terms of the originality of generated creative ideas. However, only when inter-individual differences in regular exercising were considered.</p>
<p>In people with a higher amount of regular exercise, greater HRV increase (i.e., vagal activation) during the creative ideation task was correlated with greater originality of the produced ideas, which is in accordance with some relevant research [<xref ref-type="bibr" rid="pone.0220205.ref040">40</xref>,<xref ref-type="bibr" rid="pone.0220205.ref041">41</xref>]. By contrast, in more sedentary people this association was reversed. Greater task-related HRV decrease (i.e., greater vagal withdrawal) was associated with better task performance in terms of originality, which is in line with other studies in the literature [<xref ref-type="bibr" rid="pone.0220205.ref035">35</xref>]. Importantly, regular exercising as such was not correlated with task performance, that is, on average, people exercising both more or less achieved similar levels of originality [<xref ref-type="bibr" rid="pone.0220205.ref052">52</xref>,<xref ref-type="bibr" rid="pone.0220205.ref089">89</xref>]. This null finding might take place because of study-design divergences. Contrary to former studies [<xref ref-type="bibr" rid="pone.0220205.ref051">51</xref>,<xref ref-type="bibr" rid="pone.0220205.ref053">53</xref>], we used a self-report measure of regular physical exercise [<xref ref-type="bibr" rid="pone.0220205.ref052">52</xref>]. Furthermore, while Gondola and Tuckman [<xref ref-type="bibr" rid="pone.0220205.ref053">53</xref>] reported increases in the fluency parameter of the AU task and Latorre Román et al. [<xref ref-type="bibr" rid="pone.0220205.ref051">51</xref>] found a positive effect for a composite score of multiple creative thinking tasks, we operationalized creativity by means of originality of ideas.</p>
<p>Nevertheless, the observed interaction effect suggests that greater originality in a creativity task is not only achieved by the investment of more mental effort [<xref ref-type="bibr" rid="pone.0220205.ref024">24</xref>,<xref ref-type="bibr" rid="pone.0220205.ref035">35</xref>], and more spontaneous and associative modes of thinking [<xref ref-type="bibr" rid="pone.0220205.ref032">32</xref>], but that both can lead to success, depending, among others, on the physical constitution of the person. This finding may in part explain the heterogeneity of previous results on associations between cardiac vagal control and creative performance [<xref ref-type="bibr" rid="pone.0220205.ref024">24</xref>,<xref ref-type="bibr" rid="pone.0220205.ref035">35</xref>,<xref ref-type="bibr" rid="pone.0220205.ref040">40</xref>].</p>
<p>Furthermore, the observed moderation effect at least in part supports assumptions of the cross-stressor adaptation theory [<xref ref-type="bibr" rid="pone.0220205.ref057">57</xref>], according to which changing cardiac response patterns to physical stressors by physical exercising may translate to altered modes of responding of regulation functions when confronted with psychological stressors [<xref ref-type="bibr" rid="pone.0220205.ref046">46</xref>,<xref ref-type="bibr" rid="pone.0220205.ref059">59</xref>,<xref ref-type="bibr" rid="pone.0220205.ref060">60</xref>]. People who generate more original ideas showed smaller vagal withdrawal (or even increases in heart rate variability) during task performance, when they were regularly exercising as compared to being more sedentary. This is in line with Park et al. [<xref ref-type="bibr" rid="pone.0220205.ref090">90</xref>], who reported task-related increases of HRV during a cognitive task in participants with higher flexible regulation functions (i.e., higher tonic vagal control) and decreases in participants with lower regulation functions (for short review see [<xref ref-type="bibr" rid="pone.0220205.ref015">15</xref>]).</p>
<p>The present findings need to be discussed in light of some limitations. First, the amount of regular physical exercise was only assessed by self-report and not by a behavioral index. Although the current findings need to be replicated with objective measures, similar self-reports of exercise have proven suitable to reveal reliable and meaningful associations with challenge-related cardiac responses [<xref ref-type="bibr" rid="pone.0220205.ref059">59</xref>]. Second, it might be proposed that HRV is influenced by verbal responses during the creative ideation task [<xref ref-type="bibr" rid="pone.0220205.ref091">91</xref>]. However, even if the vocalization of ideas has an impact on HRV, this cannot explain the reported associations between HRV and originality (for people exercising more or less, respectively), since the number of generated ideas (within the 6 minutes activation period) was virtually independent of task performance (originality). This is further in line with the result, that the respiration frequency (EDR) had no significant impact on the findings of this study. Third, although the eye closed baseline condition is the most frequently used approach in psychophysiological research [<xref ref-type="bibr" rid="pone.0220205.ref015">15</xref>], an additional baseline using an eyes open condition might have been valuable to evaluate the specificity of reported task-related HRV changes. Fourth, due to the cross-sectional, correlational nature of the present study, no cause-effect relationship can be deduced from the results.</p>
<p>Despite of these limitations, the findings of this research suggests that participants exercising more often in their daily life achieve higher originality with more flexible, associative, and spontaneous modes of thinking, while participants exhibiting a more sedentary lifestyle more strongly invest mental effort to reach the same quality level of produced ideas. Since solving open and ill-defined problems constitutes a daily mental challenge for many people such mechanisms might impose health risks and lower well-being in people with a sedentary lifestyle in the long run. Conversely, grounding on the cross-stressor adaptation theory the findings of the present research supports the beneficial effect of regular exercising on mental challenges in daily life [<xref ref-type="bibr" rid="pone.0220205.ref046">46</xref>,<xref ref-type="bibr" rid="pone.0220205.ref057">57</xref>].</p>
<sec id="sec013">
<title>Conclusions</title>
<p>The current study provides evidence that creative performance depends on individual differences in task-related changes in vagal activity, when taking regular exercising into account. This supports the conclusions that firstly, regular physical exercising has an impact on creativity [<xref ref-type="bibr" rid="pone.0220205.ref051">51</xref>,<xref ref-type="bibr" rid="pone.0220205.ref053">53</xref>] and secondly, higher creativity can be achieved by different strategies such as spontaneous modes of thinking and mental effort [<xref ref-type="bibr" rid="pone.0220205.ref024">24</xref>,<xref ref-type="bibr" rid="pone.0220205.ref032">32</xref>].</p>
</sec>
</sec>
<sec id="sec014">
<title>Supporting information</title>
<supplementary-material id="pone.0220205.s001" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet" position="float" xlink:href="info:doi/10.1371/journal.pone.0220205.s001" xlink:type="simple">
<label>S1 Dataset</label>
<caption>
<title/>
<p>(XLSX)</p>
</caption>
</supplementary-material>
</sec>
</body>
<back>
<ref-list>
<title>References</title>
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