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To date, the only species with complete connectomes are worms\n                  <jats:sup>1-3<\/jats:sup>\n                  and sea squirts4 (10\n                  <jats:sup>3<\/jats:sup>\n                  -10\n                  <jats:sup>4<\/jats:sup>\n                  synapses). By contrast, the fruit fly is more complex (10\n                  <jats:sup>8<\/jats:sup>\n                  synaptic connections), with a brain that supports learning and spatial memory\n                  <jats:sup>5,6<\/jats:sup>\n                  and an intricate ventral nerve cord analogous to the vertebrate spinal cord\n                  <jats:sup>7-11<\/jats:sup>\n                  . Here we report the first densely reconstructed adult fly connectome that unites the brain and ventral nerve cord, and we leverage this resource to investigate principles of neural control. We show that effector neurons (motor neurons, endocrine cells and efferent neurons targeting the viscera) are primarily influenced by sensory neurons in the same body part, forming local feedback loops. These local loops are linked by long-range circuits involving ascending and descending neurons organized into behavior-centric modules. Single ascending and descending neurons are often positioned to influence the voluntary movements of multiple body parts, together with the endocrine cells or visceral organs that support those movements. Brain regions involved in learning and navigation supervise these circuits. These results reveal an architecture that is distributed, parallelized and embodied, reminiscent of distributed control architectures in engineered systems\n                  <jats:sup>12,13<\/jats:sup>\n                  .\n                <\/jats:p>","DOI":"10.1101\/2025.07.31.667571","type":"posted-content","created":{"date-parts":[[2025,8,1]],"date-time":"2025-08-01T15:50:21Z","timestamp":1754063421000},"source":"Crossref","is-referenced-by-count":46,"title":["Distributed control circuits across a brain-and-cord connectome"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1195-0445","authenticated-orcid":false,"given":"Alexander Shakeel","family":"Bates","sequence":"first","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA; Centre for Neural Circuit and Behaviour, University of Oxford, Oxford, UK;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8033-258X","authenticated-orcid":false,"given":"Jasper S","family":"Phelps","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA; Present address: Neuroengineering Laboratory, Brain Mind Institute and Institute of Bioengineering, EPFL, Lausanne, Switzerland;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0074-6684","authenticated-orcid":false,"given":"Minsu","family":"Kim","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA; Present address: Department of Molecular and Cellular Biology, Harvard University, Cambridge, MA, USA; Present address: Center for Brain Science, Harvard University, Cambridge, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5140-9664","authenticated-orcid":false,"given":"Helen H","family":"Yang","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-4490-2028","authenticated-orcid":false,"given":"Arie","family":"Matsliah","sequence":"additional","affiliation":[{"name":"Princeton Neuroscience Institute, Princeton University, Princeton, NJ, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4411-4005","authenticated-orcid":false,"given":"Zaki","family":"Ajabi","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5542-1302","authenticated-orcid":false,"given":"Eric","family":"Perlman","sequence":"additional","affiliation":[{"name":"Yikes LLC, Baltimore, MD, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-7504-0826","authenticated-orcid":false,"given":"Kevin M","family":"Delgado","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8606-6518","authenticated-orcid":false,"given":"Mohammed Abdal Monium","family":"Osman","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5732-8117","authenticated-orcid":false,"given":"Christopher K","family":"Salmon","sequence":"additional","affiliation":[{"name":"Princeton Neuroscience Institute, Princeton University, Princeton, NJ, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jay","family":"Gager","sequence":"additional","affiliation":[{"name":"Princeton Neuroscience Institute, Princeton University, Princeton, NJ, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-7518-3465","authenticated-orcid":false,"given":"Benjamin","family":"Silverman","sequence":"additional","affiliation":[{"name":"Princeton Neuroscience Institute, Princeton University, Princeton, NJ, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0128-3458","authenticated-orcid":false,"given":"Sophia","family":"Renauld","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-2937-807X","authenticated-orcid":false,"given":"Farzaan","family":"Salman","sequence":"additional","affiliation":[{"name":"Department of Biology, West Virginia University, Morgantown, WV, USA; Present address: Department of Neurobiology, Harvard Medical School, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8140-9347","authenticated-orcid":false,"given":"Janki","family":"Patel","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3933-529X","authenticated-orcid":false,"given":"Matthew F","family":"Collie","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7860-4558","authenticated-orcid":false,"given":"Jingxuan","family":"Fan","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1638-2376","authenticated-orcid":false,"given":"Diego A","family":"Pacheco","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-3021-2389","authenticated-orcid":false,"given":"Yunzhi","family":"Zhao","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0132-9794","authenticated-orcid":false,"given":"Wenyi","family":"Zhang","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Harvard Medical School, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9167-0285","authenticated-orcid":false,"given":"Laia","family":"Serratosa Capdevilla","sequence":"additional","affiliation":[{"name":"Aelysia LTD, Bristol, UK;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2216-1465","authenticated-orcid":false,"given":"Ruairi JV","family":"Roberts","sequence":"additional","affiliation":[{"name":"Aelysia LTD, Bristol, UK;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9284-9463","authenticated-orcid":false,"given":"Eva J","family":"Munnelly","sequence":"additional","affiliation":[{"name":"Aelysia LTD, Bristol, UK;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-7769-7933","authenticated-orcid":false,"given":"Nina","family":"Griggs","sequence":"additional","affiliation":[{"name":"Aelysia LTD, Bristol, UK;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1418-2306","authenticated-orcid":false,"given":"Helen","family":"Langley","sequence":"additional","affiliation":[{"name":"Aelysia LTD, Bristol, UK;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-4560-2976","authenticated-orcid":false,"given":"Borja","family":"Moya-Llamas","sequence":"additional","affiliation":[{"name":"Aelysia LTD, Bristol, UK;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-6679-4396","authenticated-orcid":false,"given":"Zuoyu","family":"Zhang","sequence":"additional","affiliation":[{"name":"Beijing Institute for Brain Research, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China; Chinese Institute for Brain Research, Beijing, Beijing, China;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6111-7822","authenticated-orcid":false,"given":"Ryan T","family":"Maloney","sequence":"additional","affiliation":[{"name":"Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA, USA; Center for Brain Science, Harvard University, Cambridge, MA, USA; Present address: Psychology Department, Colorado College, Colorado Springs, CO, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Szi-chieh","family":"Yu","sequence":"additional","affiliation":[{"name":"Princeton Neuroscience Institute, Princeton University, Princeton, NJ, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amy R","family":"Sterling","sequence":"additional","affiliation":[{"name":"Princeton Neuroscience Institute, Princeton University, Princeton, NJ, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-7636-0746","authenticated-orcid":false,"given":"Marissa","family":"Sorek","sequence":"additional","affiliation":[{"name":"Princeton Neuroscience Institute, Princeton University, Princeton, NJ, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-9305-7042","authenticated-orcid":false,"given":"Krzysztof","family":"Kruk","sequence":"additional","affiliation":[{"name":"Eyewire, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-7752-6715","authenticated-orcid":false,"given":"Nikitas","family":"Serafetinidis","sequence":"additional","affiliation":[{"name":"Eyewire, Boston, MA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2872-1948","authenticated-orcid":false,"given":"Serene","family":"Dhawan","sequence":"additional","affiliation":[{"name":"Princeton Neuroscience Institute, Princeton University, Princeton, NJ, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-3800-3244","authenticated-orcid":false,"given":"Finja","family":"Klemm","sequence":"additional","affiliation":[{"name":"Genetics Department, Leipzig University, Leipzig, Germany;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5998-6378","authenticated-orcid":false,"given":"Paul","family":"Brooks","sequence":"additional","affiliation":[{"name":"Zoology Department, University of Cambridge, Cambridge, UK;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7929-0503","authenticated-orcid":false,"given":"Ellen","family":"Lesser","sequence":"additional","affiliation":[{"name":"Department of Molecular Genetics and Cell Biology, The University of Chicago, Chicago, IL, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3638-255X","authenticated-orcid":false,"given":"Jessica M","family":"Jones","sequence":"additional","affiliation":[{"name":"Department of Neurobiology and Biophysics, University of Washington, Seattle, WA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-4589-761X","authenticated-orcid":false,"given":"Sara E","family":"Pierce-Lundgren","sequence":"additional","affiliation":[{"name":"Department of Neurobiology and Biophysics, University of Washington, Seattle, WA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5160-9930","authenticated-orcid":false,"given":"Su-Yee","family":"Lee","sequence":"additional","affiliation":[{"name":"Department of Neurobiology and Biophysics, University of Washington, Seattle, WA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2265-1117","authenticated-orcid":false,"given":"Yichen","family":"Luo","sequence":"additional","affiliation":[{"name":"Department of Neurobiology and Biophysics, University of Washington, Seattle, WA, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4349-2880","authenticated-orcid":false,"given":"Andrew P","family":"Cook","sequence":"additional","affiliation":[{"name":"Department of Biology, West Virginia University, Morgantown, WV, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8501-6487","authenticated-orcid":false,"given":"Theresa H","family":"McKim","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Nevada Reno, Reno, NV, USA;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-8602-2308","authenticated-orcid":false,"given":"Dimitrios Stasi","family":"Giakoumas","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Florida, Gainesville, FL, USA; 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