Distributed control circuits across a brain-and-cord connectome

Author information

Author notes

Jasper S. Phelps

Present address: Neuroengineering Laboratory, Brain Mind Institute and Institute of Bioengineering, EPFL, Lausanne, Switzerland

Minsu Kim

Present address: Department of Molecular and Cellular Biology, Harvard University, Cambridge, MA, USA

Minsu Kim

Present address: Center for Brain Science, Harvard University, Cambridge, MA, USA

Farzaan Salman

Present address: Department of Neurobiology, Harvard Medical School, Boston, MA, USA

Ryan T. Maloney

Present address: Psychology Department, Colorado College, Colorado Springs, CO, USA

Dimitrios Stasi Giakoumas

Present address: Interdisciplinary Graduate Program in Neuroscience, University of Iowa, Iowa City, IA, USA

These authors contributed equally: Alexander S. Bates, Jasper S. Phelps, Minsu Kim, Helen H. Yang

Authors and Affiliations

Department of Neurobiology, Harvard Medical School, Boston, MA, USA

Alexander S. Bates, Jasper S. Phelps, Minsu Kim, Helen H. Yang, Zaki Ajabi, Kevin M. Delgado, Mohammed Abdal Monium Osman, Sophia Renauld, Janki Patel, Matthew F. Collie, Jingxuan Fan, Diego A. Pacheco, Yunzhi Zhao, Wenyi Zhang, Jasper S. Phelps, Minsu Kim, Emily Ye, Jan Drugowitsch, Rachel I. Wilson & Wei-Chung Allen Lee

Centre for Neural Circuit and Behaviour, University of Oxford, Oxford, UK

Alexander S. Bates

Princeton Neuroscience Institute, Princeton University, Princeton, NJ, USA

Arie Matsliah, Christopher K. Salmon, Jay Gager, Benjamin Silverman, Szi-chieh Yu, Amy R. Sterling, Marissa Sorek, Serene Dhawan, Austin Burke, James Hebditch, Kyle P. Willie, Ryan Willie, Emily Behnke, Sandeep Kumar, Anna Verbe, Zijin Huang, H. Sebastian Seung & Mala Murthy

Yikes LLC, Baltimore, MD, USA

Eric Perlman

Department of Biology, West Virginia University, Morgantown, WV, USA

Farzaan Salman, Farzaan Salman & Kevin C. Daly

Aelysia LTD, Bristol, UK

Laia Serratosa Capdevila, Ruairí J. V. Roberts, Eva J. Munnelly, Nina Griggs, Helen Langley & Borja Moya-Llamas

Beijing Institute for Brain Research, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing, China

Zuoyu Zhang & Tatsuo S. Okubo

Chinese Institute for Brain Research, Beijing, Beijing, China

Zuoyu Zhang & Tatsuo S. Okubo

Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA, USA

Ryan T. Maloney, Ryan T. Maloney, Benjamin L. de Bivort & Benjamin L. de Bivort

Center for Brain Science, Harvard University, Cambridge, MA, USA

Ryan T. Maloney, Ryan T. Maloney, Benjamin L. de Bivort & Benjamin L. de Bivort

Eyewire, Boston, MA, USA

Krzysztof Kruk, Nikitas Serafetinidis, Doug Bland, Anne Kristiansen, Jaime Skelton, Tom Stocks, Anthony Hernandez, Devon Jones & Celia David

Genetics Department, Leipzig University, Leipzig, Germany

Finja Klemm & Katharina Eichler

Zoology Department, University of Cambridge, Cambridge, UK

Paul Brooks

Department of Molecular Genetics and Cell Biology, The University of Chicago, Chicago, IL, USA

Ellen Lesser

Department of Neurobiology and Biophysics, University of Washington, Seattle, WA, USA

Jessica M. Jones, Sara E. Pierce-Lundgren, Su-Yee Lee, Yichen Luo, John C. Tuthill & Anthony Azevedo

School of Neuroscience, Virginia Tech, Blacksburg, VA, USA

Andrew P. Cook & Sweta Agrawal

Department of Biology, University of Nevada Reno, Reno, NV, USA

Theresa H. McKim

Department of Biology, University of Florida, Gainesville, FL, USA

Dimitrios Stasi Giakoumas, Dimitrios Stasi Giakoumas & Zepeng Yao

Molecular, Cellular, and Developmental Biology, University of California, Santa Barbara, Santa Barbara, CA, USA

Benjamin Gorko & Dustin Garner

HHMI Janelia, Ashburn, VA, USA

Justin Ellis-Joyce, Diane-Yayra Adjavon & Jan Funke

Department of Neuroscience, Johns Hopkins University, Baltimore, MD, USA

Justin Ellis-Joyce

Institute of Neuroscience, Center for Excellence in Brain Science and Intelligence Technology, Chinese Academy of Sciences, Shanghai, China

Jiayi Zhang & Fei Wang

Department of Biological Sciences, Vanderbilt University, Nashville, TN, USA

Emily C. Kophs & Marie P. Suver

Max Planck Institute for Biological Intelligence, Martinsried, Germany

Tjalda Falt & Lisa M. Fenk

Institute of Neurobiology, University of Puerto Rico Medical Sciences Campus, San Juan, Puerto Rico

Alexa M. Negron-Morales, Gabriel A. Nieves-Sanabria, Omaris Y. De Pablo-Crespo, Fernando J. Figueroa Santiago, Andrew M. Seeds & Stefanie Hampel

Zetta AI LLC, Sherrill, NY, USA

Sergiy Popovych, Nico Kemnitz, Dodam Ih, Kisuk Lee, Ran Lu, Akhilesh Halageri, J. Alexander Bae & Thomas Macrina

School of Informatics, University of Edinburgh, Edinburgh, UK

Ben Jourdan

Japan Advanced Institute of Science and Technology (JAIST), Nomi, Japan

Gregory Schwartzman

Molecular Brain Physiology and Behavior, LIMES Institute, University of Bonn, Bonn, Germany

Damian D. Demarest & Michael J. Pankratz

Department of Neuroscience, West Virginia University, Morgantown, WV, USA

Kevin C. Daly

Allen Institute for Brain Science, Seattle, WA, USA

Sven Dorkenwald & Forrest Collman

Florida Chemical Senses Institute, University of Florida, Gainesville, FL, USA

Zepeng Yao

McKnight Brain Institute, University of Florida, Gainesville, FL, USA

Zepeng Yao

Genetics Institute, University of Florida, Gainesville, FL, USA

Zepeng Yao

Department of Neuroscience, Zuckerman Mind Brain Behavior Institute, Columbia University, New York, NY, USA

Stephen J. Huston

Neurobiology Division, MRC Laboratory of Molecular Biology, Cambridge, UK

Tomke Stürner & Gregory S. X. E. Jefferis

Department of Biochemistry and Molecular Biology, University of Nevada Reno, Reno, NV, USA

Meet Zandawala

Neurobiology and Genetics, Theodor-Boveri-Institute, Biocenter, Julius-Maximilians-University of Würzburg, Am Hubland, Würzburg, Germany

Meet Zandawala

Computer Science Department, Princeton University, Princeton, NJ, USA

H. Sebastian Seung

F.M. Kirby Neurobiology Center, Boston Children’s Hospital, Harvard Medical School, Boston, MA, USA

Wei-Chung Allen Lee

Champalimaud Neuroscience Programme, Champalimaud Centre for the Unknown, Lisbon, Portugal

Sofia Pinto

Department of Biology, Leipzig University, Leipzig, Germany

Wolf Huetteroth

RWTH Aachen University, Aachen, Germany

Zequan Liu

Authors

Alexander S. Bates

Jasper S. Phelps

Minsu Kim

Helen H. Yang

Arie Matsliah

Zaki Ajabi

Eric Perlman

Kevin M. Delgado

Mohammed Abdal Monium Osman

Christopher K. Salmon

Jay Gager

Benjamin Silverman

Sophia Renauld

Farzaan Salman

Janki Patel

Matthew F. Collie

Jingxuan Fan

Diego A. Pacheco

Yunzhi Zhao

Wenyi Zhang

Laia Serratosa Capdevila

Ruairí J. V. Roberts

Eva J. Munnelly

Nina Griggs

Helen Langley

Borja Moya-Llamas

Zuoyu Zhang

Ryan T. Maloney

Szi-chieh Yu

Amy R. Sterling

Marissa Sorek

Krzysztof Kruk

Nikitas Serafetinidis

Serene Dhawan

Finja Klemm

Paul Brooks

Ellen Lesser

Jessica M. Jones

Sara E. Pierce-Lundgren

Su-Yee Lee

Yichen Luo

Andrew P. Cook

Theresa H. McKim

Dimitrios Stasi Giakoumas

Benjamin Gorko

Justin Ellis-Joyce

Jiayi Zhang

Emily C. Kophs

Tjalda Falt

Alexa M. Negron-Morales

Austin Burke

James Hebditch

Kyle P. Willie

Ryan Willie

Sergiy Popovych

Nico Kemnitz

Dodam Ih

Kisuk Lee

Ran Lu

Akhilesh Halageri

J. Alexander Bae

Ben Jourdan

Gregory Schwartzman

Damian D. Demarest

Emily Behnke

Doug Bland

Anne Kristiansen

Jaime Skelton

Tom Stocks

Dustin Garner

Anthony Hernandez

Sandeep Kumar

Kevin C. Daly

Sven Dorkenwald

Forrest Collman

Marie P. Suver

Lisa M. Fenk

Michael J. Pankratz

Zepeng Yao

Fei Wang

Stephen J. Huston

Tomke Stürner

Gregory S. X. E. Jefferis

Katharina Eichler

Andrew M. Seeds

Stefanie Hampel

Sweta Agrawal

Tatsuo S. Okubo

Meet Zandawala

Thomas Macrina

Diane-Yayra Adjavon

Jan Funke

John C. Tuthill

Anthony Azevedo

H. Sebastian Seung

Benjamin L. de Bivort

Mala Murthy

Jan Drugowitsch

Rachel I. Wilson

Wei-Chung Allen Lee

Consortia

The BANC-FlyWire Consortium

Alexander S. Bates

, Jasper S. Phelps

, Minsu Kim

, Helen H. Yang

, Arie Matsliah

, Zaki Ajabi

, Eric Perlman

, Kevin M. Delgado

, Mohammed Abdal Monium Osman

, Christopher K. Salmon

, Jay Gager

, Benjamin Silverman

, Sophia Renauld

, Farzaan Salman

, Janki Patel

, Matthew F. Collie

, Jingxuan Fan

, Diego A. Pacheco

, Yunzhi Zhao

, Wenyi Zhang

, Laia Serratosa Capdevila

, Ruairí J. V. Roberts

, Eva J. Munnelly

, Nina Griggs

, Helen Langley

, Borja Moya-Llamas

, Zuoyu Zhang

, Ryan T. Maloney

, Szi-chieh Yu

, Amy R. Sterling

, Marissa Sorek

, Krzysztof Kruk

, Nikitas Serafetinidis

, Serene Dhawan

, Finja Klemm

, Paul Brooks

, Ellen Lesser

, Jessica M. Jones

, Sara E. Pierce-Lundgren

, Su-Yee Lee

, Yichen Luo

, Andrew P. Cook

, Theresa H. McKim

, Dimitrios Stasi Giakoumas

, Benjamin Gorko

, Justin Ellis-Joyce

, Jiayi Zhang

, Emily C. Kophs

, Tjalda Falt

, Alexa M. Negron-Morales

, Austin Burke

, James Hebditch

, Kyle P. Willie

, Ryan Willie

, Sergiy Popovych

, Nico Kemnitz

, Dodam Ih

, Kisuk Lee

, Ran Lu

, Akhilesh Halageri

, J. Alexander Bae

, Ben Jourdan

, Gregory Schwartzman

, Damian D. Demarest

, Emily Behnke

, Doug Bland

, Anne Kristiansen

, Jaime Skelton

, Tom Stocks

, Dustin Garner

, Anthony Hernandez

, Sandeep Kumar

, Kevin C. Daly

, Sven Dorkenwald

, Forrest Collman

, Marie P. Suver

, Lisa M. Fenk

, Michael J. Pankratz

, Zepeng Yao

, Fei Wang

, Stephen J. Huston

, Tomke Stürner

, Gregory S. X. E. Jefferis

, Katharina Eichler

, Andrew M. Seeds

, Stefanie Hampel

, Sweta Agrawal

, Tatsuo S. Okubo

, Meet Zandawala

, Thomas Macrina

, Diane-Yayra Adjavon

, Jan Funke

, John C. Tuthill

, Anthony Azevedo

, H. Sebastian Seung

, Benjamin L. de Bivort

, Mala Murthy

, Jan Drugowitsch

, Rachel I. Wilson

, Wei-Chung Allen Lee

, Anna Verbe

, Gabriel A. Nieves-Sanabria

, Devon Jones

, Zijin Huang

, Sofia Pinto

, Celia David

, Omaris Y. De Pablo-Crespo

, Emily Ye

, Wolf Huetteroth

, Zequan Liu

& Fernando J. Figueroa Santiago

Corresponding authors

Correspondence to

Alexander S. Bates, Jasper S. Phelps, Mala Murthy, Jan Drugowitsch, Rachel I. Wilson or Wei-Chung Allen Lee.

Supplementary information

Supplementary Information (download PDF )

Detailed description of the columns of each of the Supplementary Data files.

Reporting Summary (download PDF )

Supplementary Data 1 (download TXT )

Metadata categories and terms. Table of categories of annotations applied to BANC neurons and the list of terms used in each category.

Supplementary Data 2 (download TXT )

BANC neuron metadata. Per-neuron metadata for the proofread/roughly proofread neurons in our BANC release, with cross-dataset match identifiers to FAFB, MANC, hemibrain, FANC and maleCNS where assigned. This is the reference table for every BANC analysis in the paper.

Supplementary Data 3 (download TXT )

Updated FAFB-FlyWire neuron metadata. Per-neuron metadata for the FAFB-FlyWire v783 neurons used in BANC's cross-dataset comparisons. We re-annotated several columns (super_class, cell_class, cell_sub_class, body_part_*, cell_function) to follow BANC conventions; cell_type names are unchanged from the FAFB project.

Supplementary Data 4 (download TXT )

Updated MANC neuron metadata. Per-neuron metadata for the 49,052 VNC neurons from MANC v1.2.1, with super_class / cell_class / cell_sub_class / body_part_* / cell_function aligned to BANC conventions. cell_type names are unchanged from the MANC project.

Supplementary Data 5 (download TXT )

maleCNS neuron metadata. Per-neuron metadata for the neurons in the male CNS v0.9 release. Cell typing is all from the maleCNS project, but annotations have been aligned to the conventions of the BANC, to help users compare and co-analyze these datasets.

Supplementary Data 6 (download TXT )

ANs and DNs with PCA-UMAP coordinates and cluster assignments. Per-neuron metadata for the ANs and DNs included in the connectivity-PCA-UMAP analysis (Fig. 3d, Extended Data Fig. 6), with cluster assignments.

Supplementary Data 7 (download TXT )

Effector neurons with UMAP coordinates and functional cluster assignments. Per-neuron metadata for the efferent neurons (motor + visceral/circulatory) in the connectivity-UMAP analysis (Extended Data Fig. 4f), with cluster assignments.

Supplementary Data 8 (download TXT )

CNS network analysis with spectral clustering and UMAP embedding. Per-neuron metadata for the 50,568 intrinsic neurons that entered the CNS-network spectral clustering analysis (Fig. 6a, Extended Data Fig. 10). Spectral clustering parameters: min connection strength = 1, cluster count = 13, cluster seed = 10, embedding seed = 3 (banc_888).

Supplementary Data 9 (download TXT )

Literature review on cell function for ascending, descending and visual projection neurons. Curated list of literature-validated functional roles used to interpret the AN/DN clusters and visual-projection influences

Supplementary Data 10 (download TXT )

Bounding boxes for known dataset artifacts that negatively impact neuronal reconstruction. 22 bounding boxes delineating regions with known data-quality issues in the BANC dataset. Coordinates are in BANC raw-voxel space (1 voxel = 4 × 4 × 45 nm).

Peer Review File (download PDF )

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Cite this article

Bates, A.S., Phelps, J.S., Kim, M. et al. Distributed control circuits across a brain-and-cord connectome.

Nature (2026). https://doi.org/10.1038/s41586-026-10735-w

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Received: 01 August 2025

Accepted: 29 May 2026

Published: 08 June 2026

DOI: https://doi.org/10.1038/s41586-026-10735-w