The supplied source is a bibliography covering how nervous systems generate and adapt movement, with a strong focus on insects—especially Drosophila—and on the use of connectomes, computational models and neuromechanical simulations. The references span foundational work on central pattern generators and rhythmic activity, studies of locomotion and proprioception, recent reconstructions of the fly nervous system, and efforts to connect neural structure with behaviour.
From rhythmic circuits to motor control
Several references examine the neural circuits that generate rhythmic actions such as walking, swimming, flying, breathing and heartbeat. Early work includes T. G. Brown’s 1914 analysis of rhythmic activity and C. S. Sherrington’s The Integrative Action of the Nervous System (1920). Later studies explored swimmeret coordination in crayfish, flight control in locusts, locomotion in leeches, and the small but complex stomatogastric nervous system.
Research on central pattern generators has also addressed mammalian locomotion, spinal motor networks, neuronal redundancy and the interaction between feedforward and feedback control. More recent work describes rotational neural dynamics in spinal circuits, sparse firing in hybrid spinal models, and the emergence of coordinated locomotor activity from cell-type-specific connectivity.
Connectomes and the Drosophila nervous system
A substantial portion of the bibliography concerns connectomics: the reconstruction and analysis of neural wiring. Studies have reported connectomes of the male and female Drosophila ventral nerve cord, the adult central brain, and the male central nervous system. Other work examines sexual dimorphism, ascending and descending neurons, synaptic architecture in leg and wing premotor networks, and quantitative neuroanatomy.
The references also emphasize that a connectome alone does not fully explain brain function. Researchers have developed computational models constrained by connectomic data, including models of sensorimotor processing, visual-system activity, social behaviour and head-direction circuits. Tools such as neuPrint and the Connectome Annotation Versioning Engine (CAVE) support access to and analysis of these datasets.
Locomotion, proprioception and behaviour
The cited studies investigate how animals adapt movement to their environment and internal state. Topics include leg proprioception in Drosophila, the recruitment of leg motor neurons, presynaptic inhibition of proprioceptive signals, adaptive walking on miniature and split-belt treadmills, and neural mechanisms that halt movement in specific contexts.
Other references examine grooming, including descending neurons that coordinate anterior grooming and dedicated command circuits for grooming control. Work on descending networks describes how command signals are transformed into population-level motor control, while recent preprints address dedicated brain circuits for forward walking and the distinction between central and peripheral control of coordinated walking.
Models linking brains, bodies and behaviour
The bibliography includes neuromechanical and physics-based approaches to animal movement. NeuroMechFly is described as a neuromechanical model of adult Drosophila melanogaster, while whole-body physics simulations model fruit-fly locomotion. Related work includes a virtual rodent designed to predict neural activity across behaviours and a preprint asking whether a worm brain could control a fly body.
Computational methods cited in the source include neural differential equations, attractor-based models, canonical circuits for phase-amplitude coupling, conductance-based neuron databases, and mathematical and theoretical approaches to neural systems. The list also identifies software for neural modelling, pose estimation and behavioural tracking, including JAX, DeepLabCut, Anipose and FicTrac.
References
- Büschges, A. & Ache, J. M. “Motor control on the move: from insights in insects to general mechanisms.” Physiol. Rev. 105, 975–1031 (2025). Article
- Gosgnach, S. “The mammalian locomotor CPG: revealing the contents of the black box.” J. Neurophysiol. 133, 472–478 (2025). Article
- Takemura, S.-Y. et al. “A connectome of the male Drosophila ventral nerve cord.” Preprint (2024).
- Azevedo, A. et al. “Connectomic reconstruction of a female Drosophila ventral nerve cord.” Nature 631, 360–368 (2024). Article
- 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). Article
- 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 (1960). Article
- Wilson, D. M. “The central nervous control of flight in a locust.” J. Exp. Biol. 38, 471–490 (1961). Article
- Daur, N., Nadim, F. & Bucher, D. “The complexity of small circuits: the stomatogastric nervous system.” Curr. Opin. Neurobiol. 41, 1–7 (2016). Article
- Friesen, W. O. & Kristan, W. B. “Leech locomotion: swimming, crawling, and decisions.” Curr. Opin. Neurobiol. 17, 704–711 (2007). Article
- Calabrese, R. L., Norris, B. J. & Wenning, A. “The neural control of heartbeat in invertebrates.” Curr. Opin. Neurobiol. 41, 68–77 (2016). Article
- Del Negro, C. A., Funk, G. D. & Feldman, J. L. “Breathing matters.” Nat. Rev. Neurosci. 19, 351–367 (2018). Article
- Shepherd, G. M., Grillner, S. & Katz, P. S. In Handbook of Brain Microcircuits, 561–568 (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 (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). Article
- 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 (2024). Article
- Holmes, P., Full, R. J., Koditschek, D. & Guckenheimer, J. “The dynamics of legged locomotion: models, analyses, and challenges.” SIAM Rev. 48, 207–304 (2006). Article
- 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 (2023). Article
- 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). Article
- Marin, E. C. et al. “Systematic annotation of a complete adult male Drosophila nerve cord connectome reveals principles of functional organisation.” Preprint (2024).
- Bargmann, C. I. & Marder, E. “From the connectome to brain function.” Nat. Methods 10, 483–490 (2013). Article
- 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 (2021). Article
- Shiu, P. K. et al. “A Drosophila computational brain model reveals sensorimotor processing.” Nature 634, 210–219 (2024). Article
- 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). Article
- Cowley, B. R. et al. “Mapping model units to visual neurons reveals population code for social behaviour.” Nature 629, 1100–1108 (2024). Article
- 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). Article
- Agrawal, S. et al. “Central processing of leg proprioception in Drosophila.” eLife 9, e60299 (2020).
- Azevedo, A. W. et al. “A size principle for recruitment of Drosophila leg motor neurons.” eLife 9, e56754 (2020). Article
- 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). Article
- Gouwens, N. W. & Wilson, R. I. “Signal propagation in Drosophila central neurons.” J. Neurosci. 29, 6239–6249 (2009). Article
- 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). Article
- Scheffer, L. K. et al. “A connectome and analysis of the adult Drosophila central brain.” eLife 9, e57443 (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). Article
- 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). Article
- Guo, L., Zhang, N. & Simpson, J. H. “Descending neurons coordinate anterior grooming behavior in Drosophila.” Curr. Biol. 32, 823–833 (2022). Article
- Hampel, S., Franconville, R., Simpson, J. H. & Seeds, A. M. “A neural command circuit for grooming movement control.” eLife 4, e08758 (2015).
- Briggman, K. L. & Kristan, W. B. “Multifunctional pattern-generating circuits.” Ann. Rev. Neurosci. 31, 271–294 (2008). Article
- Scheffer, L. K. “Synapse detection efficiency in EM Drosophila connectomics.” Preprint at bioRxiv (2025).
- Sapkal, N. et al. “Central versus peripheral neural control of a coordinated walking pattern in Drosophila.” Preprint at bioRxiv (2026).
- Lobato-Rios, V. et al. “NeuroMechFly, a neuromechanical model of adult Drosophila melanogaster.” Nat. Methods 19, 620–627 (2022). Article
- Vaxenburg, R. et al. “Whole-body physics simulation of fruit fly locomotion.” Nature 643, 1312–1320 (2025). Article
- Braun, J., Hurtak, F., Wang-Chen, S. & Ramdya, P. “Descending networks transform command signals into population motor control.” Nature 630, 686–694 (2024). Article
- Dallmann, C. J. et al. “A dedicated brain circuit controls forward walking in Drosophila.” Preprint at bioRxiv (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). Article
- Alvarez, J. L., Morrison, K. & Curto, C. “Attractor-Based Models for Sequences and Pattern Generation in Neural Circuits.” Neur. Comput. 38, 257–291 (2026). Article
- 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 (2003). Article
- Elowitz, M. B. & Leibler, S. “A synthetic oscillatory network of transcriptional regulators.” Nature 403, 335–338 (2000). Article
- Stricker, J. et al. “A fast, robust and tunable synthetic gene oscillator.” Nature 456, 516–519 (2008). Article
- Marder, E., Kedia, S. & Morozova, E. O. “New insights from small rhythmic circuits.” Curr. Opin. Neurobiol. 76, 102610 (2022). Article
- Aldarondo, D. et al. “A virtual rodent predicts the structure of neural activity across behaviours.” Nature 632, 594–602 (2024). Article
- Brunton, B. W., Abe, E. T., Hu, L. J. & Tuthill, J. C. “The digital sphinx: can a worm brain control a fly body?” Preprint (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). Article
- 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 (2016).
- Bradbury, J. et al. “JAX: composable transformations of Python+NumPy programs v.0.3.13.” GitHub (2018).
- Kidger, P. “On neural differential equations.” Preprint (2022).
- Zung, J. L. et al. “An updated catalogue of split-GAL4 driver lines for descending neurons in Drosophila melanogaster.” Preprint (2025).
- Mathis, A. et al. “DeepLabCut: markerless pose estimation of user-defined body parts with deep learning.” Nat. Neurosci. 21, 1281–1289 (2018).
- Karashchuk, L. et al. “Anipose: a toolkit for robust markerless 3D pose estimation.” Cell Rep. 36, 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 (2014).
- Pratt, B. G. et al. “Proprioceptive limit detectors contribute to sensorimotor control of the Drosophila leg.” Nat. Commun. 17, 2664 (2026). Article
Source: www.nature.com


