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Here, using optogenetic stimulation of various brainstem, thalamic, and dopaminergic cell populations in mice trained on an action-timing task, we manipulated actions preceding the timing decision to causally test these hypotheses. Our findings show that bidirectional regulation of actions during the waiting period leads to corresponding temporal shifts in timing, with a consistent quantitative relationship between changes in the number of waiting-period actions and shifts in stop timing. Notably, nigrostriatal dopamine encodes actions, but not timing, directly, influencing timing only when waiting-period actions are modulated. These findings reveal a behavioral integration mechanism underlying action timing, in which actions performed during the waiting period function as a pacemaker for the internal clock.<\/jats:p>","DOI":"10.1126\/sciadv.aea5558","type":"journal-article","created":{"date-parts":[[2026,1,2]],"date-time":"2026-01-02T18:59:44Z","timestamp":1767380384000},"update-policy":"https:\/\/doi.org\/10.34133\/aaas_crossmark","source":"Crossref","is-referenced-by-count":2,"title":["A behavioral integration mechanism underlies action timing"],"prefix":"10.1126","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0009-0008-5171-1714","authenticated-orcid":true,"given":"Patrick","family":"Strassmann","sequence":"first","affiliation":[{"name":"Molecular Neurobiology Laboratory, The Salk Institute for Biological Studies, La Jolla, CA 92037, USA."},{"name":"Neurosciences Graduate Program, University of California San Diego, La Jolla, CA 92093, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5246-5796","authenticated-orcid":true,"given":"Xiaochun","family":"Cai","sequence":"additional","affiliation":[{"name":"Molecular Neurobiology Laboratory, The Salk Institute for Biological Studies, La Jolla, CA 92037, USA."},{"name":"Neurosciences Graduate Program, University of California San Diego, La Jolla, CA 92093, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-1300-0580","authenticated-orcid":true,"given":"Baibing","family":"Zhang","sequence":"additional","affiliation":[{"name":"New Cornerstone Science Laboratory, Center for Motor Control and Disease, Key Laboratory of Brain Functional Genomics, East China Normal University, Shanghai 200062, China."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4838-6323","authenticated-orcid":true,"given":"Christopher D.","family":"Howard","sequence":"additional","affiliation":[{"name":"Molecular Neurobiology Laboratory, The Salk Institute for Biological Studies, La Jolla, CA 92037, USA."},{"name":"Department of Neuroscience, Oberlin College, Oberlin, OH 44074, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jonathan","family":"Cook","sequence":"additional","affiliation":[{"name":"Molecular Neurobiology Laboratory, The Salk Institute for Biological Studies, La Jolla, CA 92037, USA."},{"name":"Neurosciences Graduate Program, University of California San Diego, La Jolla, CA 92093, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1106-4013","authenticated-orcid":true,"given":"Xin","family":"Jin","sequence":"additional","affiliation":[{"name":"Molecular Neurobiology Laboratory, The Salk Institute for Biological Studies, La Jolla, CA 92037, USA."},{"name":"New Cornerstone Science Laboratory, Center for Motor Control and Disease, Key Laboratory of Brain Functional Genomics, East China Normal University, Shanghai 200062, China."},{"name":"NYU-ECNU Institute of Brain and Cognitive Science, New York University Shanghai, Shanghai 200062, China."}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"221","reference":[{"key":"e_1_3_2_2_2","doi-asserted-by":"publisher","DOI":"10.1038\/nrn1764"},{"key":"e_1_3_2_3_2","doi-asserted-by":"publisher","DOI":"10.1038\/npp.2010.113"},{"key":"e_1_3_2_4_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.tics.2018.07.011"},{"key":"e_1_3_2_5_2","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.11386"},{"key":"e_1_3_2_6_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cobeha.2016.01.006"},{"key":"e_1_3_2_7_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2018.03.045"},{"key":"e_1_3_2_8_2","doi-asserted-by":"publisher","DOI":"10.1038\/85191"},{"key":"e_1_3_2_9_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0010324"},{"key":"e_1_3_2_10_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2017.02.029"},{"key":"e_1_3_2_11_2","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.54831"},{"key":"e_1_3_2_12_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cobeha.2016.01.003"},{"key":"e_1_3_2_13_2","first-page":"10","article-title":"Ongoing behavior predicts perceptual report of interval duration","volume":"8","author":"Gouv\u00eaa T. 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