A neural circuit architecture for angular integration in Drosophila
Open accessAbstract
Many animals keep track of their angular heading over time while navigating through their environment. However, a neural-circuit architecture for computing heading has not been experimentally defined in any species. Here we describe a set of clockwise- and anticlockwise-shifting neurons in the Drosophila central complex whose wiring and physiology provide a means to rotate an angular heading estimate based on the fly's angular velocity. We show that each class of shifting neurons exists in two subtypes, with spatiotemporal activity profiles that suggest different roles for each subtype at the start and end of tethered-walking turns. Shifting neurons are required for the heading system to properly track the fly's heading in the dark, and stimulation of these neurons induces predictable shifts in the heading signal. The central features of this biological circuit are analogous to those of computational models proposed for head-direction cells in rodents and may shed light on how neural systems, in general, perform integration.
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No statistical methods were used to predetermine sample sizes, but the sample sizes are similar to those reported in previous publications.
“No statistical methods were used to predetermine sample sizes but our sample sizes are similar to those reported in previous publications23,24,33,34,37,38.”
From Multimodal cue integration and learning in a neural representation of head direction (2025) · cites this paper as [38]
The source paper’s Methods, under Data analysis (page unavailable), says its sample size was not chosen using a statistical method. It does not say that its sample sizes were similar to those in previous publications, nor can it establish that comparison for the citing authors’ sample sizes. (LLM, conf=0.86)
Experimental — accuracy not yet measuredCompartments of PEN2 neurons in the protocerebral bridge respond entirely differently from their compartments in the ellipsoid body.
“Similarly, compartments of PEN2 neurons in the protocerebral bridge have been shown to respond entirely differently from their compartments in the ellipsoid body (Green et al., 2017; Turner-Evans et al., 2019).”
From A connectome and analysis of the adult Drosophila central brain (2020) · cites this paper as Green et al., 2017
In “Spatiotemporal relationships across cells” (page unavailable), the paper reports that P-EN2 signals projected from the bridge predicted a broad valley, whereas imaging found a sharp peak in the ellipsoid body. But “entirely differently” overstates the contrast: the paper says the ellipsoid-body activity retained a turn-related asymmetry consistent with the bridge activity. (LLM, conf=0.85)
Experimental — accuracy not yet measuredThe EB is known to be involved in navigation, supporting persistent activity correlated to heading direction.
“Another well studied neuropil, the EB, is known to be involved in navigation, supporting persistent activity correlated to heading direction57–59.”
From Network statistics of the whole-brain connectome of Drosophila (2024) · cites this paper as [58]
There is a single bump of activity in the EPG ensemble whose angular velocity generally correlates with the fly's angular velocity.
“we found that there is a single bump of activity in the EPG ensemble whose angular velocity generally correlates with the fly’s angular velocity (Green et al., 2017; Seelig and Jayaraman, 2015; Turner-Evans et al., 2017).”
From Neural circuit mechanisms for steering control in walking Drosophila (2025) · cites this paper as Green et al., 2017
The central complex is the primary locus of spatial computations in arthropods.
“The central complex is the primary locus of spatial computations in arthropods.”
From Transforming representations of movement from body- to world-centric space (2022) · cites this paper as [19]
World-centric heading direction θ is computed in the ellipsoid body and sent to the protocerebral bridge, while body-centric translation direction φ is relayed to the nodulus.
“Here, world-centric heading direction (θ) is computed in the ellipsoid body and sent to the protocerebral bridge (PB)17–19 while body-centric translation direction (φ) is relayed to the nodulus (NO)20.”
From Transforming representations of movement from body- to world-centric space (2022) · cites this paper as [19]
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“…another possibility that could explain the bump’s movements is raised by a parallel study, which provides further evidence for P-ENs serving a role in angular inte- gration and describes a second subtype of P-EN neurons that likely also influences the position of the E-PG bump (Green et al., 2017).”
ResultAttractor and integrator networks in the brain
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“In a striking example of convergent evolution [155, 151], Drosophila compute HD estimates using apparently very similar dynamics [156, 157, 153, 152].”
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BackgroundDistinct Dopamine Receptor Pathways Underlie the Temporal Sensitivity of Associative Learning
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InfluentialTransforming representations of movement from body- to world-centric space
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“EB projection neurons (EPG neurons or “compass neurons”) send this heading information to the protocerebral bridge (PB) 7,8 (Fig.”
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“For example, the two PEN neurons below have very similar morphology but quite distinct inputs, as shown in Fig 7 (PEN1 and PEN2 neurons, in fact, have been shown to have different functional activity in [34]).”
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BackgroundMethodInfluentialNetwork statistics of the whole-brain connectome of Drosophila
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BackgroundInfluentialTwo brain pathways initiate distinct forward walking programs in Drosophila
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Functional Imaging and Optogenetics in Drosophila
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BackgroundMethodConverting an allocentric goal into an egocentric steering signal
Mussells Pires, P., Zhang, L., Parache, V., Abbott, L. F., & Maimon, G. (2024) · Nature · doi:10.1038/s41586-023-07006-3 · Cited by 110 · see citing works
“For both two-photon calcium imaging and patch-clamp experiments, we placed flies in a virtual reality setup described previously 34 .”
BackgroundMethodInfluentialSelf-citationNeural dynamics and architecture of the heading direction circuit in zebrafish
Petrucco, L., Lavian, H., Wu, Y. K., Svara, F., Štih, V., & Portugues, R. (2023) · Nature Neuroscience · doi:10.1038/s41593-023-01308-5 · Cited by 110 · see citing works
A decentralised neural model explaining optimal integration of navigational strategies in insects
Sun, X., Yue, S., & Mangan, M. (2020) · eLife · doi:10.7554/elife.54026 · Cited by 96 · see citing works
“Insects track their orientation through head-direction cells Seelig and Jayaraman, 2015 whose concurrent firing pattern forms a single bump of activity that shifts around the ring as the animal turns (measured through local visual [Green et al., 2017; Turner-Evans et al., 2017], global visual (Heinze and Homberg, 2007…”
The head direction circuit of two insect species
Pisokas, I., Heinze, S., & Webb, B. (2020) · eLife · doi:10.7554/elife.53985 · Cited by 84 · see citing works
Decoding locomotion from population neural activity in moving C. elegans
Hallinen, K. M., Dempsey, R., Scholz, M., Yu, X., Linder, A. N., Randi, F., Sharma, A. K., Shaevitz, J. W., & Leifer, A. M. (2021) · eLife · doi:10.7554/elife.66135 · Cited by 106 · see citing works
“For example, population recordings from the central complex in Drosophila reveal that the animal’s heading is represented in the population by a bump of neural activity in a ring attractor network (Kim et al., 2017; Green et al., 2017).”
Savelli, F., & Knierim, J. J. (2019) · Journal of Experimental Biology · doi:10.1242/jeb.188912 · Cited by 93 · see citing works
“Vice versa, a process conceptually and computationally equivalent to path integration can be applied to a purely directional reference frame – based on the integration of angular velocity to update orientation irrespective of position – as occurs in the nervous system (Green et al., 2017; Knierim and Zhang, 2012; Skag…”
BackgroundMultisensory coding of angular head velocity in the retrosplenial cortex
Keshavarzi, S., Bracey, E. F., Faville, R., Campagner, D., Tyson, A. L., Lenzi, S. C., Branco, T., & Margrie, T. W. (2021) · Neuron · doi:10.1016/j.neuron.2021.10.031 · Cited by 88 · see citing works
“It is widely accepted that the updating of this heading direction signal arises from angular velocity integration (Blair and Sharp, 1995; Green et al., 2017; Laurens and Angelaki, 2018; McNaughton et al., 1991; McNaughton et al., 1995; TurnerEvans et al., 2017), with theoretical models of head direction computation re…”
BackgroundToroidal topology of population activity in grid cellsPreprint
Gardner, R. J., Hermansen, E., Pachitariu, M., Burak, Y., Baas, N. A., Dunn, B. A., Moser, M., & Moser, E. I. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.02.25.432776 · Cited by 88 · see citing works
Context-dependent representations of movement in Drosophila dopaminergic reinforcement pathways
Zolin, A., Cohn, R., Pang, R., Siliciano, A. F., Fairhall, A. L., & Ruta, V. (2021) · Nature Neuroscience · doi:10.1038/s41593-021-00929-y · Cited by 81 · see citing works
Transforming a head direction signal into a goal-oriented steering command
Westeinde, E. A., Kellogg, E., Dawson, P. M., Lu, J., Hamburg, L., Midler, B., Druckmann, S., & Wilson, R. I. (2024) · Nature · doi:10.1038/s41586-024-07039-2 · Cited by 85 · see citing works
“Available data indicate that there are two complete linearized topographic maps of head direction in Δ7 cells, positioned side-by-side and formatted as two cycles of a sinusoidal function over neural space 5,24,47,56 .”
Muscles that move the retina augment compound eye vision in Drosophila
Fenk, L. M., Avritzer, S. C., Weisman, J. L., Nair, A., Randt, L., Mohren, T. L., Siwanowicz, I., & Maimon, G. (2022) · Nature · doi:10.1038/s41586-022-05317-5 · Cited by 82 · see citing works
Self-citationFlexible navigational computations in the Drosophila central complex
Fisher, Y. E. (2022) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2021.12.001 · Cited by 74 · see citing works
Neuronal constituents and putative interactions within the Drosophila Ellipsoid Body Neuropil
Omoto, J. J., Nguyen, B.-C. M., Kandimalla, P., Lovick, J. K., Donlea, J. M., & Hartenstein, V. (2018) · Frontiers in Neural Circuits · doi:10.3389/fncir.2018.00103 · Cited by 72 · see citing works
Neural circuit mechanisms for steering control in <i>walking Drosophila</i>
Rayshubskiy, A., Holtz, S. L., Bates, A. S., Vanderbeck, Q. X., Capdevila, L. S., Rockwell, V., & Wilson, R. I. (2020) · doi:10.1101/2020.04.04.024703 · Cited by 64 · see citing works
“As expected from previous work, we found that there is a single bump of activity in the EPG ensemble whose angular velocity generally correlates with the fly’s angular velocity (Green et al., 2017; Seelig and Jayaraman, 2015; Turner-Evans et al., 2017).”
Building a heading signal from anatomically defined neuron types in the Drosophila central complex
Green, J., & Maimon, G. (2018) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2018.06.010 · Cited by 79 · see citing works
Self-citationFlight-Induced Compass Representation in the Monarch Butterfly Heading Network
Beetz, M. J., Kraus, C. M., Franzke, M., Dreyer, D., Strube-Bloss, M. F., Rößler, W., Warrant, E. J., Merlin, C., & el Jundi, B. (2021) · Current Biology · doi:10.1016/j.cub.2021.11.009 · Cited by 73 · see citing works
Diverse Food-Sensing Neurons Trigger Idiothetic Local Search in Drosophila
Corfas, R. A., Sharma, T., & Dickinson, M. H. (2019) · Current Biology · doi:10.1016/j.cub.2019.03.004 · Cited by 60 · see citing works
Dopamine promotes head direction plasticity during orienting movements
Fisher, Y. E., Marquis, M., D’Alessandro, I., & Wilson, R. I. (2022) · Nature · doi:10.1038/s41586-022-05485-4 · Cited by 67 · see citing works
“35 ) or PEN1 (ref.”
InfluentialA projectome of the bumblebee central complex
Sayre, M. E., Templin, R., Chávez, J., Kempenaers, J., & Heinze, S. (2021) · eLife · doi:10.7554/elife.68911 · Cited by 75 · see citing works
The structure of behavioral variation within a genotype
Werkhoven, Z., Bravin, A., Skutt-Kakaria, K., Reimers, P., Pallares, L. F., Ayroles, J., & de Bivort, B. (2019) · bioRxiv · doi:10.7554/elife.64988 · Cited by 49 · see citing works
“…changes when we inactivated or activated neurons in the Central Complex, a cluster of neuropils involved in locomotor behaviors (Buchanan et al., 2015; Ofstad et al., 2011; Kottler et al., 2019) and heading estimation (Seelig and Nature, 2015; and Shah et al., 2017; Kakaria and de Bivort, 2017).”
BackgroundVisual processing in the fly, from photoreceptors to behavior
Currier, T. A., Pang, M. M., & Clandinin, T. R. (2023) · GENETICS · doi:10.1093/genetics/iyad064 · Cited by 63 · see citing works
The Brain Compass: A Perspective on How Self-Motion Updates the Head Direction Cell Attractor
Laurens, J., & Angelaki, D. E. (2018) · Neuron · doi:10.1016/j.neuron.2017.12.020 · Cited by 67 · see citing works
Emergent behaviour and neural dynamics in artificial agents tracking odour plumes
Singh, S. H., van Breugel, F., Rao, R. P. N., & Brunton, B. W. (2023) · Nature Machine Intelligence · doi:10.1038/s42256-022-00599-w · Cited by 61 · see citing works
Three brain states in the hippocampus and cortex
Kay, K., & Frank, L. M. (2018) · Hippocampus · doi:10.1002/hipo.22956 · Cited by 63 · see citing works
Neuroarchitecture of the dung beetle central complex
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Encoding and control of orientation to airflow by a set of Drosophila fan-shaped body neurons
Currier, T. A., Matheson, A. M. M., & Nagel, K. I. (2020) · eLife · doi:10.7554/elife.61510 · Cited by 60 · see citing works
Parallel encoding of recent visual experience and self-motion during navigation in Drosophila
Shiozaki, H. M., & Kazama, H. (2017) · Nature Neuroscience · doi:10.1038/nn.4628 · Cited by 60 · see citing works
The evolution of honey bee dance communication: a mechanistic perspective
Barron, A. B., & Plath, J. A. (2017) · Journal of Experimental Biology · doi:10.1242/jeb.142778 · Cited by 60 · see citing works
“…publications have provided some substantial experimental evidence towards an additional integration of orientation angles and angular velocity: Green et al. (2017) and Turner-Evans et al. (2017) have identified a type of neuron (tile neuron) which connects the EB to the PB. Activity in these…”
BackgroundA DNAzyme-powered cross-catalytic circuit for amplified intracellular imaging
Zou, L., Wu, Q. D., Zhou, Y., Gong, X. Q., Liu, X., & Wang, F. (2019) · Chemical Communications · doi:10.1039/c9cc02637a · Cited by 60 · see citing works
Brain-wide representations of ongoing behavior: a universal principle?
Kaplan, H. S., & Zimmer, M. (2020) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2020.02.008 · Cited by 57 · see citing works
A brainstem integrator for self-location memory and positional homeostasis in zebrafish
Yang, E., Zwart, M., James, B., Rubinov, M., Wei, Z., Narayan, S., Vladimirov, N., Mensh, B. D., Fitzgerald, J. E., & Ahrens, M. B. (2022) · Cell · doi:10.1016/j.cell.2022.11.022 · Cited by 61 · see citing works
Mapping the neural dynamics of locomotion across the Drosophila brain
Brezovec, B. E., Berger, A. B., Hao, Y. A., Chen, F., Druckmann, S., & Clandinin, T. R. (2024) · Current Biology · doi:10.1016/j.cub.2023.12.063 · Cited by 58 · see citing works
Functional network topography of the medial entorhinal cortex
Obenhaus, H. A., Zong, W., Jacobsen, R. I., Rose, T., Donato, F., Chen, L., Cheng, H., Bonhoeffer, T., Moser, M., & Moser, E. I. (2022) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2121655119 · Cited by 55 · see citing works
“One of the best illustrations of this concept in non-mammalian species is the discovery of a ring-shaped head-direction (HD) circuit in the centroid complex of fruit flies (Green et al., 2017; Kim et al., 2017; Seelig and Jayaraman, 2015), whose anatomical layout and wiring mirrors the predictions of ring attractor ne…”
BackgroundSchilling, M., & Cruse, H. (2020) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1007804 · Cited by 55 · see citing works
Connectomics and the neural basis of behaviour
Galili, D. S., Jefferis, G. S. X. E., & Costa, M. (2022) · Current Opinion in Insect Science · doi:10.1016/j.cois.2022.100968 · Cited by 54 · see citing works
“This circuit relies on combining both internal self-motion cues (angular velocity) [55,56] and external cues (polarised light, visual landmarks, mechanosensory wind cues or others), carried by ring neurons.”
BackgroundHulse, B. K., Haberkern, H., Franconville, R., Turner-Evans, D. B., Takemura, S., Wolff, T., Noorman, M., Dreher, M., Dan, C., Parekh, R., et al. (2020) · doi:10.1101/2020.12.08.413955 · Cited by 392 · see citing works
“Within the PB, FB columnar neurons inherit a head direction bump directly from EPG neurons and indirectly through Δ7 neurons (Figure 68Bi) (Franconville et al., 2018; Green et al., 2017; Turner-Evans et al., 2017; Turner-Evans et al., 2020).”
BackgroundResultInfluentialZittrell, F., Pfeiffer, K., & Homberg, U. (2020) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2005192117 · Cited by 49 · see citing works
“heading signals are represented in the CX in a compass-like, topographic fashion (4, 8, 16, 28, 29).”
BackgroundInfluentialA Multi-regional Network Encoding Heading and Steering Maneuvers in Drosophila
Shiozaki, H., Ohta, K., & Kazama, H. (2020) · Neuron · doi:10.1016/j.neuron.2020.01.009 · Cited by 50 · see citing works
Karagyozov, D., Skanata, M. M., Lesar, A., & Gershow, M. H. (2018) · Cell Reports · doi:10.1016/j.celrep.2018.10.013 · Cited by 49 · see citing works
Gkanias, E., Risse, B., Mangan, M., & Webb, B. H. (2019) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1007123 · Cited by 49 · see citing works
“5 ̊ degrees apart [33], whereas the fly’s compass covers at least 270 ̊, changing by around� 35 ̊ per column [34] and might be assumed to form a 360 ̊ representation of space [32].”
BackgroundFiring patterns in a fractional-order FithzHugh–Nagumo neuron model
Yao, Z., Sun, K., & He, S. (2022) · Nonlinear Dynamics · doi:10.1007/s11071-022-07690-2 · Cited by 48 · see citing works
The brain behind straight-line orientation in dung beetles
el Jundi, B., Baird, E., Byrne, M. J., & Dacke, M. (2019) · Journal of Experimental Biology · doi:10.1242/jeb.192450 · Cited by 48 · see citing works
“Feature detection and orientation tuning in the Drosophila central complex.”
BackgroundInfluentialKottler, B., Faville, R., Bridi, J. C., & Hirth, F. (2019) · Current Biology · doi:10.1016/j.cub.2019.01.017 · Cited by 47 · see citing works
“, using the same Gal4 driver line, revealed that a turn is performed when the activity of a single PB glomerulus takes over the rest of the population, leading to locally restricted neuronal activity [19, 44].”
InfluentialA rise-to-threshold process for a relative-value decision
Vijayan, V., Wang, F., Wang, K., Chakravorty, A., Adachi, A., Akhlaghpour, H., Dickson, B. J., & Maimon, G. (2023) · Nature · doi:10.1038/s41586-023-06271-6 · Cited by 49 · see citing works
Xie, X., Tabuchi, M., Brown, M. P., Mitchell, S. P., Wu, M. N., & Kolodkin, A. L. (2017) · eLife · doi:10.7554/elife.25328 · Cited by 46 · see citing works
“DOI: 10.7554/eLife.25328 2 of 33 fly’s head direction coordinates (Seelig and Jayaraman, 2015; Turner-Evans et al., 2017; Green et al., 2017).”
BackgroundInfluentialSullivan, L. F., Warren, T. L., & Doe, C. Q. (2019) · eLife · doi:10.7554/elife.43482 · Cited by 46 · see citing works
Madhav, M. S., & Cowan, N. J. (2020) · Annual Review of Control Robotics and Autonomous Systems · doi:10.1146/annurev-control-060117-104856 · Cited by 45 · see citing works
“An apparatus that allows two-photon calcium imaging while head-fixed flies walk on an air-supported ball (145) enabled the discovery of a neural correlate of the hypothesized ring-attractor angular integration system for encoding orientation in the central complex of flies (146, 147).”
BackgroundConnectomic reconstruction predicts visual features used for navigation
Garner, D., Kind, E., Lai, J. Y. H., Nern, A., Zhao, A., Houghton, L., Sancer, G., Wolff, T., Rubin, G. M., Wernet, M. F., et al. (2024) · Nature · doi:10.1038/s41586-024-07967-z · Cited by 44 · see citing works
“1a), where they together form a complex recurrent neural network 5,41,42 (Fig.”
Self-Organized Attractor Dynamics in the Developing Head Direction Circuit
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Automated real-time quantification of group locomotor activity in Drosophila melanogaster
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The brain of a nocturnal migratory insect, the Australian Bogong moth
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Olfactory navigation in arthropods
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Vision for navigation: What can we learn from ants?
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The neuronal building blocks of the navigational toolkit in the central complex of insects
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Two Compasses in the Central Complex of the Locust Brain
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“This range is close to the twofold representation of 360° space of E-PG neurons (equivalents to locust CL1 neurons) in the fly (Green et al., 2017), which was also a necessary assumption for a computational model of path integration in bees (Stone et al.”
BackgroundInfluentialQuantifying behavior to solve sensorimotor transformations: advances from worms and flies
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“Employing such a setup has already revealed many of the sensorimotor processes that underlie spatial navigation in Drosophila [63,64].”
BackgroundRecent Advances in the Genetic Dissection of Neural Circuits in Drosophila
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Neural networks for navigation: From connections to computations
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The cognitive (lateral) hypothalamus
Sharpe, M. J. (2023) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2023.08.019 · Cited by 38 · see citing works
- Authors
- Green, J., Adachi, A., Shah, K. K., Hirokawa, J. D., Magani, P. S., & Maimon, G.
- Affiliations
- Rockefeller University (US)
- Corresponding author
- Maimon, Gaby
- Venue
- Nature
- Published
- 2017-05-22
- Volume
- 546
- Issue
- 7656
- Pages
- 101-106
- ISSN
- 0028-0836 (ISSN-L)
- 1476-4687
- PMC
- PMC6320684
- OpenAlex
- W2614915054
- Cited by
- 331
- References
- 43
- Citations by year
- 2026: 24 · 2025: 41 · 2024: 39 · 2023: 39 · 2022: 37 · 2021: 30 · 2020: 31 · 2019: 41 · 2018: 31 · 2017: 17
- FWCI
- 16.20
- Type
- journal article
- Language
- en
- Open access
- green
- europepmc.org
- Licence
- www.springer.com/tdm (tdm)
- Locations
- Nature (publishedVersion)
- PubMed (publishedVersion)
- Europe PMC (PubMed Central) (acceptedVersion)
- PubMed Central (acceptedVersion)
- RePEc: Research Papers in Economics (submittedVersion)
- Funders
- National Institute for Health and Care Research
- National Institutes of Health (P40OD018537)
- National Institute on Drug Abuse (DP2DA035148)
- National Institute of Neurological Disorders and Stroke (R01NS104934)
- Topics
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurobiology and Insect Physiology Research
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Life Sciences › Neuroscience › Neurology › Vestibular and auditory disorders
- Keywords
- central complex, angular integration, Drosophila, spatial navigation, persistent activity, turning behavior, neural circuit architecture, angular velocity
- MeSH terms
- Animals, Darkness, Drosophila melanogaster, Female, Flight, Animal, Models, Neurological, Neural Pathways, Neurons, Orientation, Rotation, Space Perception, Walking, Spatio-Temporal Analysis
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