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However, the mechanisms by which electric fields modulate cortical network activity are only partially understood. Our aim was to determine the modulation of spontaneous cortical activity by static electric fields and their underlying network mechanisms, which we investigated\n                      <jats:italic>in vitro<\/jats:italic>\n                      and\n                      <jats:italic>in silico<\/jats:italic>\n                      . Neocortical slices were exposed to constant fields of varying intensities and directions. We measured their effect on slow oscillations (SOs, &lt;1 Hz), which consist of Up (active) and Down (silent) states, and on the high\u2010frequency content (\u03b2: 15\u201330 Hz, \u03b3: 30\u201390 Hz) during Up states. We found that DC fields ranging from \u22126 to +6 V\/m induced an exponential increase in the frequency of SOs through the regulation of neuronal excitability and the duration of Down states, while hardly affecting Up state duration. A computational model based on the mean\u2010field theory of attractor dynamics provided a mechanistic and quantitative description of the network dynamics underlying such precise modulation of slow oscillatory frequency. The modulation of high frequencies by positive DC fields was less consistent, the high\u2010frequency power varying with the intensity of the fields only in a fraction of slices. Negative DC fields of increasing intensities progressively and effectively reversed the kainate\u2010induced high\u2010frequency power. DC fields precisely modulate emergent cortical network activity even through millivolt\u2010scale effects on individual neurons. This effect is specific for different parameters of cortical oscillations. We discuss the underlying mechanisms and implications.\n                      <jats:boxed-text content-type=\"graphic\" position=\"anchor\">\n                        <jats:graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" mimetype=\"image\/png\" position=\"anchor\" specific-use=\"enlarged-web-image\" xlink:href=\"graphic\/tjp70380-gra-0001-m.png\">\n                          <jats:alt-text>image<\/jats:alt-text>\n                        <\/jats:graphic>\n                      <\/jats:boxed-text>\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Key points<\/jats:title>\n                    <jats:p>\n                      <jats:list list-type=\"bullet\">\n                        <jats:list-item>\n                          <jats:p>\n                            We studied the impact that direct current (DC) electric fields had on the slow (&lt;1 Hz) and fast (\u03b2: 15\u201330 Hz and \u03b3: 30\u201390 Hz) frequencies spontaneously generated by the cerebral cortex\n                            <jats:italic>in vitro<\/jats:italic>\n                            .\n                          <\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>We found that weak (&lt;6 V\/m) DC fields can control the frequency of slow oscillations generated by the cortical network, and we explored the underlying mechanisms both experimentally and in a computer model.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>We conclude that, despite producing only millivolt\u2010scale effects at the single\u2010neuron level, DC fields can robustly modulate cortical population activity, with efficacy varying across frequency bands.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>These findings are relevant for the design of effective tDCS protocols for therapeutical purposes.<\/jats:p>\n                        <\/jats:list-item>\n                      <\/jats:list>\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1113\/jp277400","type":"journal-article","created":{"date-parts":[[2026,2,12]],"date-time":"2026-02-12T06:35:01Z","timestamp":1770878101000},"page":"2131-2151","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Modulation of slow and fast oscillations by direct current stimulation in the cerebral cortex\n                    <i>in vitro<\/i>"],"prefix":"10.1113","volume":"604","author":[{"given":"Mattia","family":"D'Andola","sequence":"first","affiliation":[{"name":"Systems Neuroscience Institut d'Investigacions Biom\u00e8diques August Pi i Sunyer (IDIBAPS)  Barcelona Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Julia F.","family":"Weinert","sequence":"additional","affiliation":[{"name":"Systems Neuroscience Institut d'Investigacions Biom\u00e8diques August Pi i Sunyer (IDIBAPS)  Barcelona Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Joana","family":"Covelo","sequence":"additional","affiliation":[{"name":"Systems Neuroscience Institut d'Investigacions Biom\u00e8diques August Pi i Sunyer (IDIBAPS)  Barcelona Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maurizio","family":"Mattia","sequence":"additional","affiliation":[{"name":"Natl. 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