Büschges, A. & Ache, J. M. Motor control on the move: from insights in insects to general mechanisms. Physiol. Rev. 105, 975–1031 https://doi.org/10.1152/physrev.00009.2024 (2025).
Gosgnach, S. The mammalian locomotor CPG: revealing the contents of the black box. J. Neurophysiol. 133, 472–478 https://doi.org/10.1152/jn.00238.2024 (2025).
Takemura, S.-Y. et al. A connectome of the male Drosophila ventral nerve cord. Preprint at https://doi.org/10.7554/eLife.97769.1 (2024).
Azevedo, A. et al. Connectomic reconstruction of a female Drosophila ventral nerve cord. Nature 631, 360–368 (2024).
Bates, A. S. et al. Distributed control circuits across a brain-and-cord connectome. Nature 656, 957–970 (2026).
Berg, S. et al. Sexual dimorphism in the complete connectome of the Drosophila male central nervous system. Cell 189, 5504–5526.e15 (2025).
Sherrington, C. S. The Integrative Action of the Nervous System (Yale Univ. Press, 1920).
Brown, T. G. On the nature of the fundamental activity of the nervous centres; together with an analysis of the conditioning of rhythmic activity in progression, and a theory of the evolution of function in the nervous system. J. Physiol. 48, 18–46 (1914).
Hughes, G. M. & Wiersma, C. A. G. The co-ordination of swimmeret movements in the crayfish, Procambarus Clarkii (Girard). J. Exp. Biol. 37, 657–670 https://doi.org/10.1242/jeb.37.4.657 (1960).
Wilson, D. M. The central nervous control of flight in a locust. J. Exp. Biol. 38, 471–490 https://doi.org/10.1242/jeb.38.2.471 (1961).
Daur, N., Nadim, F. & Bucher, D. The complexity of small circuits: the stomatogastric nervous system. Curr. Opin. Neurobiol. 41, 1–7 (2016).
Friesen, W. O. & Kristan, W. B. Leech locomotion: swimming, crawling, and decisions. Curr. Opin. Neurobiol. 17, 704–711 (2007).
Calabrese, R. L., Norris, B. J. & Wenning, A. The neural control of heartbeat in invertebrates. Curr. Opin. Neurobiol. 41, 68–77 (2016).
Del Negro, C. A., Funk, G. D. & Feldman, J. L. Breathing matters. Nat. Rev. Neurosci. 19, 351–367 (2018).
Shepherd, G. M., Grillner, S. & Katz, P. S. in Handbook of Brain Microcircuits (eds Shepherd, G. M. & Grillner, S.) 561–568 https://doi.org/10.1093/med/9780190636111.003.0051 (Oxford Univ. Press, 2017).
Calabrese, R. L. & Marder, E. Degenerate neuronal and circuit mechanisms important for generating rhythmic motor patterns. Physiol. Rev. 105, 95–135 https://doi.org/10.1152/physrev.00003.2024 (2024).
Lindén, H., Petersen, P. C., Vestergaard, M. & Berg, R. W. Movement is governed by rotational neural dynamics in spinal motor networks. Nature 610, 526–531 (2022).
Strohmer, B., Najarro, E., Ausborn, J., Berg, R. W. & Tolu, S. Sparse firing in a hybrid central pattern generator for spinal motor circuits. Neur. Comput. 36, 759–780 https://doi.org/10.1162/neco_a_01660 (2024).
Holmes, P., Full, R. J., Koditschek, D. & Guckenheimer, J. The dynamics of legged locomotion: models, analyses, and challenges. SIAM Rev. 48, 207–304 (2006).
Ijspeert, A. J. & Daley, M. A. Integration of feedforward and feedback control in the neuromechanics of vertebrate locomotion: a review of experimental, simulation and robotic studies. J. Exp. Biol. 226, jeb245784 https://doi.org/10.1242/jeb.245784 (2023).
Lesser, E. et al. Synaptic architecture of leg and wing premotor control networks in Drosophila. Nature 631, 369–377 (2024).
Stürner, T. et al. Comparative connectomics of Drosophila descending and ascending neurons. Nature 643, 158–172 (2025).
Marin, E. C. et al. Systematic annotation of a complete adult male Drosophila nerve cord connectome reveals principles of functional organisation. Preprint at https://doi.org/10.7554/eLife.97766.1 (2024).
Bargmann, C. I. & Marder, E. From the connectome to brain function. Nat. Methods 10, 483–490 (2013).
Scheffer, L. K. & Meinertzhagen, I. A. A connectome is not enough—what is still needed to understand the brain of Drosophila? J. Exp. Biol. 224, jeb242740 https://doi.org/10.1242/jeb.242740 (2021).
Shiu, P. K. et al. A Drosophila computational brain model reveals sensorimotor processing. Nature 634, 210–219 (2024).
Kakaria, K. S. & de Bivort, B. L. Ring attractor dynamics emerge from a spiking model of the entire protocerebral bridge. Front. Behav. Neurosci. 11, 1662–5153 (2017).
Lappalainen, J. K. et al. Connectome-constrained networks predict neural activity across the fly visual system. Nature 634, 1132–1140 (2024).
Cowley, B. R. et al. Mapping model units to visual neurons reveals population code for social behaviour. Nature 629, 1100–1108 (2024).
Duan, S., Dong, L. L. & Fiete, I. From synapses to dynamics: obtaining function from structure in a connectome constrained model of the head direction circuit. In Proc 39th Conference on Neural Information Processing Systems 128293–128320 (NeurIPS, 2026).
Büschges, A. Role of local nonspiking interneurons in the generation of rhythmic motor activity in the stick insect. J. Neurobiol. 27, 488–512 (1995).
Agrawal, S. et al. Central processing of leg proprioception in Drosophila. eLife 9, e60299, https://doi.org/10.7554/eLife.60299 (2020).
Azevedo, A. W. et al. A size principle for recruitment of Drosophila leg motor neurons. eLife 9, e56754 (2020).
Liu, T. X., Davoudian, P. A., Lizbinski, K. M. & Jeanne, J. M. Connectomic features underlying diverse synaptic connection strengths and subcellular computation. Curr. Biol. 32, 559–569 (2022).
Gouwens, N. W. & Wilson, R. I. Signal propagation in Drosophila central neurons. J. Neurosci. 29, 6239–6249 (2009).
Prinz, A. A., Billimoria, C. P. & Marder, E. Alternative to hand-tuning conductance-based models: construction and analysis of databases of model neurons. J. Neurophysiol. 90, 3998–4015 (2003).
Sapkal, N. et al. Neural circuit mechanisms underlying context-specific halting in Drosophila. Nature 634, 191–200 (2024).
Scheffer, L. K. et al. A connectome and analysis of the adult Drosophila central brain. eLife 9, e57443 https://doi.org/10.7554/eLife.57443 (2020).
Buhmann, J. et al. Automatic detection of synaptic partners in a whole-brain Drosophila electron microscopy data set. Nat. Methods 18, 771–774 (2021).
Pratt, B. G., Lee, S.-Y. J., Chou, G. M. & Tuthill, J. C. Miniature linear and split-belt treadmills reveal mechanisms of adaptive motor control in walking Drosophila. Curr. Biol. 34, 4368–4381 (2024).
Dallmann, C. J. et al. Selective presynaptic inhibition of leg proprioception in behaving Drosophila. Nature 647, 445–453 (2025).
Guo, L., Zhang, N. & Simpson, J. H. Descending neurons coordinate anterior grooming behavior in Drosophila. English. Curr. Biol. 32, 823–833 (2022).
Hampel, S., Franconville, R., Simpson, J. H. & Seeds, A. M. A neural command circuit for grooming movement control. eLife 4, e08758 https://doi.org/10.7554/eLife.08758 (2015).
Briggman, K. L. & Kristan, W. B. Multifunctional pattern-generating circuits. Ann. Rev. Neurosci. 31, 271–294 https://doi.org/10.1146/annurev.neuro.31.060407.125552 (2008).
Scheffer, L. K. Synapse detection efficiency in EM Drosophila connectomics. Preprint at bioRxiv https://doi.org/10.1101/2025.10.16.682869 (2025).
Sapkal, N. et al. Central versus peripheral neural control of a coordinated walking pattern in Drosophila. Preprint at bioRxiv https://doi.org/10.64898/2026.04.29.721658 (2026).
Lobato-Rios, V. et al. NeuroMechFly, a neuromechanical model of adult Drosophila melanogaster. Nat. Methods 19, 620–627 (2022).
Vaxenburg, R. et al. Whole-body physics simulation of fruit fly locomotion. Nature 643, 1312–1320 (2025).
Braun, J., Hurtak, F., Wang-Chen, S. & Ramdya, P. Descending networks transform command signals into population motor control. Nature 630, 686–694 (2024).
Dallmann, C. J. et al. A dedicated brain circuit controls forward walking in Drosophila. Preprint at bioRxiv https://doi.org/10.64898/2026.01.04.697356 (2026).
Amari, S.-I. Mathematics of Neural Networks (Sangyo Tosho, 1978).
Onslow, A. C. E., Jones, M. W. & Bogacz, R. A canonical circuit for generating phase-amplitude coupling. PLoS ONE 9, e102591 (2014).
Alvarez, J. L., Morrison, K. & Curto, C. Attractor-Based Models for Sequences and Pattern Generation in Neural Circuits. Neur. Comput. 38, 257–291 (2026).
Wandler, F. D., Lemberger, B. K., McLean, D. L. & Murray, J. M. Coordinated spinal locomotor network dynamics emerge from cell-type-specific connectivity patterns. eLife 14, RP106658 (2025).
Dvornyk, V., Vinogradova, O. & Nevo, E. Origin and evolution of circadian clock genes in prokaryotes. Proc. Natl Acad. Sci. USA 100, 2495–2500 https://doi.org/10.1073/pnas.0130099100 (2003).
Elowitz, M. B. & Leibler, S. A synthetic oscillatory network of transcriptional regulators. Nature 403, 335–338 (2000).
Stricker, J. et al. A fast, robust and tunable synthetic gene oscillator. Nature 456, 516–519 (2008).
Marder, E., Kedia, S. & Morozova, E. O. New insights from small rhythmic circuits. Curr. Opin. Neurobiol. 76, 102610 (2022).
Aldarondo, D. et al. A virtual rodent predicts the structure of neural activity across behaviours. Nature 632, 594–602 (2024).
Brunton, B. W., Abe, E. T., Hu, L. J. & Tuthill, J. C. The digital sphinx: can a worm brain control a fly body? Preprint at https://doi.org/10.7554/eLife.111516.1 (2026).
Plaza, S. M. et al. neuPrint: An open access tool for EM connectomics. Front. Neuroinform. 16, 1662–5196 (2022).
Dorkenwald, S. et al. CAVE: Connectome Annotation Versioning Engine. Nat. Methods 22, 1112–1120 (2025).
Dayan, P. & Abbott, L. F. Theoretical Neuroscience: Computational and Mathematical Modeling of Neural Systems (MIT Press, 2005).
Schneider-Mizell, C. M. et al. Quantitative neuroanatomy for connectomics in Drosophila. eLife 5, e12059 https://doi.org/10.7554/eLife.12059 (2016).
Bradbury, J. et al. JAX: composable transformations of Python+NumPy programs v.0.3.13. GitHub http://github.com/jax-ml/jax (2018).
Kidger, P. On neural differential equations. Preprint at https://doi.org/10.48550/arXiv.2202.02435 (2022).
Zung, J. L. et al. An updated catalogue of split-GAL4 driver lines for descending neurons in Drosophila melanogaster. Preprint at https://doi.org/10.7554/eLife.107450.1 (2025).
Mathis, A. et al. DeepLabCut: markerless pose estimation of user-defined body parts with deep learning. Nat. Neurosci. 21, 1281–1289 https://doi.org/10.1038/s41593-018-0209-y (2018).
Karashchuk, L. et al. Anipose: a toolkit for robust markerless 3D pose estimation. Cell Rep. 36, 109730 https://doi.org/10.1016/j.celrep.2021.109730 (2021).
Moore, R. J. et al. FicTrac: A visual method for tracking spherical motion and generating fictive animal paths. J. Neurosci. Methods 225, 106–119 https://doi.org/10.1016/j.jneumeth.2014.01.010 (2014).
Pratt, B. G. et al. Proprioceptive limit detectors contribute to sensorimotor control of the Drosophila leg. Nat. Commun. 17, 2664 (2026).