Neural dynamics for landmark orientation and angular path integration
Open accessAbstract
Many animals navigate using a combination of visual landmarks and path integration. In mammalian brains, head direction cells integrate these two streams of information by representing an animal's heading relative to landmarks, yet maintaining their directional tuning in darkness based on self-motion cues. Here we use two-photon calcium imaging in head-fixed Drosophila melanogaster walking on a ball in a virtual reality arena to demonstrate that landmark-based orientation and angular path integration are combined in the population responses of neurons whose dendrites tile the ellipsoid body, a toroidal structure in the centre of the fly brain. The neural population encodes the fly's azimuth relative to its environment, tracking visual landmarks when available and relying on self-motion cues in darkness. When both visual and self-motion cues are absent, a representation of the animal's orientation is maintained in this network through persistent activity, a potential substrate for short-term memory. Several features of the population dynamics of these neurons and their circular anatomical arrangement are suggestive of ring attractors, network structures that have been proposed to support the function of navigational brain circuits.
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Processing within the central complex enables an insect to establish and hold a heading relative to an external navigational reference.
“Processing within the central complex enables an insect to establish and hold a heading relative to an external navigational reference [45,51].”
From Phenomenal interface theory: a model for basal consciousness (2025) · cites this paper as [51]
The Abstract and Discussion report that central-complex neurons in walking flies encode orientation relative to visual landmarks and maintain a compass-like representation when landmarks are absent. This supports establishing and retaining an internal heading reference, but the study does not show that this processing enables the insect to hold a behavioral heading; the Discussion describes that navigational role as a possibility. (LLM, conf=0.84)
Experimental — accuracy not yet measuredThe head direction of the insect with respect to external cues is tracked by the activity of E-PG neurons in the ellipsoid body.
“The head direction of the insect with respect to external cues is tracked by the activity of a set of EB neurons known as E-PGs [13,14], also colloquially referred to as ‘compass neurons’.”
From A model of cue integration as vector summation in the insect brain (2023) · cites this paper as [14]
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 [34]
Experimental — accuracy not yet measuredP{R60D05-Gal4}attP2 drives Gal4 expression in EPG neurons.
“P{R60D05-Gal4}attP2 drives Gal4 expression in EPG neurons, as reported previously34,41, and its construction was described previously61.”
From Multimodal cue integration and learning in a neural representation of head direction (2025) · cites this paper as [34]
The 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 [57]
The neural representation of a heading can be maintained even if the fix on the external reference is temporarily lost, enabling insects to maintain a heading and to store vectors of travelled paths in memory.
“The neural representation of that heading can be maintained even if the fix on the external reference is temporarily lost [51,52], which enables insects to maintain a heading and to store vectors of travelled paths in memory [53].”
From Phenomenal interface theory: a model for basal consciousness (2025) · cites this paper as [51]
Head direction neurons (EPG neurons) can influence steering via central complex neurons that compare head direction signals with internal goal signals.
“Head direction neurons (EPG neurons) (Seelig and Jayaraman, 2015) can influence steering (Giraldo et al., 2018; Green et al., 2019) via central complex neurons that compare head direction signals with internal goal signals (Beetz et al., 2022; Martin et al., 2015; Mussells Pires et al., 2024; Shiozaki et al., 2020; Westeinde et al., 2024).”
From Neural circuit mechanisms for steering control in walking Drosophila (2025) · cites this paper as Seelig and Jayaraman, 2015
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 Seelig and Jayaraman, 2015
Both speed and direction parameters are under feedback control, as sensory reafference continuously informs the brain about the body's instantaneous forward velocity and heading.
“Both parameters are also under feedback control, as sensory reafference continuously informs the brain about the body’s instantaneous forward velocity76–78 and heading79”
From Fine-grained descending control of steering in walking Drosophila (2024) · cites this paper as [79]
The central complex is the brain region that compares the animal's current heading to an internal goal direction.
“the brain region that compares the animal’s current heading to an internal goal direction62,63,76,77.”
From Fine-grained descending control of steering in walking <i>Drosophila</i> (2023) · cites this paper as [76]
Both speed and direction parameters are under feedback control, as sensory reafference continuously informs the brain about the body's instantaneous forward velocity and heading.
“Both parameters are also under feedback control, as sensory reafference continuously informs the brain about the body’s instantaneous forward velocity73–75 and heading76.”
From Fine-grained descending control of steering in walking <i>Drosophila</i> (2023) · cites this paper as [76]
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 [17]
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 [17]
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Cites
Ultrasensitive fluorescent proteins for imaging neuronal activity
Chen, T.-W., Wardill, T. J., Sun, Y., Pulver, S. R., Renninger, S. L., Baohan, A., Schreiter, E. R., Kerr, R. A., Orger, M. B., Jayaraman, V., et al. (2013) · Nature · doi:10.1038/nature12354 · Cited by 6,474 · see citing works
Self-citationShifts in Selective Visual Attention: Towards the Underlying Neural Circuitry
Christof Koch, A., & Ullman, S. (1987) · Springer eBooks · doi:10.1007/978-94-009-3833-5_5 · Cited by 3,739 · see citing works
<b>CircStat</b> : A <i>MATLAB</i> Toolbox for Circular Statistics
Berens, P. (2009) · Journal of Statistical Software · doi:10.18637/jss.v031.i10 · Cited by 3,099 · see citing works
Taube, J., Muller, R., & Ranck, J. (1990) · The Journal of Neuroscience · doi:10.1523/jneurosci.10-02-00420.1990 · Cited by 1,809 · see citing works
A GAL4-Driver Line Resource for Drosophila Neurobiology
Jenett, A., Rubin, G. M., Ngo, T.-T. B., Shepherd, D., Murphy, C., Dionne, H., Pfeiffer, B. D., Cavallaro, A., Hall, D., Jeter, J., et al. (2012) · Cell Reports · doi:10.1016/j.celrep.2012.09.011 · Cited by 1,449 · see citing works
ScanImage: Flexible software for operating laser scanning microscopes
Pologruto, T. A., Sabatini, B. L., & Svoboda, K. (2003) · BioMedical Engineering OnLine · doi:10.1186/1475-925x-2-13 · Cited by 1,297 · see citing works
The Head Direction Signal: Origins and Sensory-Motor Integration
Taube, J. S. (2007) · Annual Review of Neuroscience · doi:10.1146/annurev.neuro.29.051605.112854 · Cited by 916 · see citing works
Homing by path integration in a mammal
Mittelstaedt, M.-L., & Mittelstaedt, H. (1980) · Die Naturwissenschaften · doi:10.1007/bf00450672 · Cited by 768 · see citing works
Neurocomputational models of working memory
Durstewitz, D., Seamans, J. K., & B, T. J. S. A. (2000) · Nature Neuroscience · doi:10.1038/81460 · Cited by 710 · see citing works
Desert ant navigation: how miniature brains solve complex tasks
Wehner, R. (2003) · Journal of Comparative Physiology A · doi:10.1007/s00359-003-0431-1 · Cited by 577 · see citing works
The central complex and the genetic dissection of locomotor behaviour
Strauß, R. (2002) · Current Opinion in Neurobiology · doi:10.1016/s0959-4388(02)00385-9 · Cited by 429 · see citing works
Active flight increases the gain of visual motion processing in Drosophila
Maimon, G., Straw, A. D., & Dickinson, M. H. (2010) · Nature Neuroscience · doi:10.1038/nn.2492 · Cited by 420 · see citing works
Distinct memory traces for two visual features in the Drosophila brain
Liu, G., Seiler, H., Wen, A., Zars, T. D., Ito, K., Wolf, R., Heisenberg, M., & Liu, L. (2006) · Nature · doi:10.1038/nature04381 · Cited by 407 · see citing works
Visual place learning in Drosophila melanogaster
Ofstad, T. A., Zuker, C. S., & Reiser, M. B. (2011) · Nature · doi:10.1038/nature10131 · Cited by 399 · see citing works
Organization and Functional Roles of the Central Complex in the Insect Brain
Pfeiffer, K., & Homberg, U. (2014) · Annual Review of Entomology · doi:10.1146/annurev-ento-011613-162031 · Cited by 392 · see citing works
Analysis of a spatial orientation memory in Drosophila
Neuser, K., Triphan, T., Mronz, M., Poeck, B., & Strauß, R. (2008) · Nature · doi:10.1038/nature07003 · Cited by 363 · see citing works
Maplike Representation of Celestial E -Vector Orientations in the Brain of an Insect
Heinze, S., & Homberg, U. (2007) · Science · doi:10.1126/science.1135531 · Cited by 360 · see citing works
Mizumori, S., & Williams, J. (1993) · Journal of Neuroscience · doi:10.1523/jneurosci.13-09-04015.1993 · Cited by 359 · see citing works
Deep homology of arthropod central complex and vertebrate basal ganglia.
Strausfeld, N. J., & Hirth, F. (2013) · Science · doi:10.1126/science.1231828 · Cited by 354 · see citing works
Neuronal architecture of the central complex in Drosophila melanogaster
Hanesch, U., Fischbach, K., & Heisenberg, M. (2004) · Cell and Tissue Research · doi:10.1007/bf00261838 · Cited by 349 · see citing works
Two-photon calcium imaging from head-fixed Drosophila during optomotor walking behavior
Seelig, J. D., Chiappe, M. E., Lott, G. K., Dutta, A., Osborne, J. E., Reiser, M. B., & Jayaraman, V. (2010) · Nature Methods · doi:10.1038/nmeth.1468 · Cited by 345 · see citing works
Self-citationPersistent neural activity: prevalence and mechanisms
Major, G., & Tank, D. (2004) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2004.10.017 · Cited by 320 · see citing works
Feature detection and orientation tuning in the Drosophila central complex
Seelig, J. D., & Jayaraman, V. (2013) · Nature · doi:10.1038/nature12601 · Cited by 290 · see citing works
Self-citationWolff, T., Iyer, N. A., & Rubin, G. M. (2015) · Journal of Comparative Neurology · doi:10.1002/cne.23705 · Cited by 286 · see citing works
Internally organized mechanisms of the head direction sense
Peyrache, A., Lacroix, M. M., Petersen, P. C., & Buzsáki, G. (2015) · Nature Neuroscience · doi:10.1038/nn.3968 · Cited by 274 · see citing works
A modular display system for insect behavioral neuroscience
Reiser, M. B., & Dickinson, M. H. (2007) · Journal of Neuroscience Methods · doi:10.1016/j.jneumeth.2007.07.019 · Cited by 272 · see citing works
Sun Compass Integration of Skylight Cues in Migratory Monarch Butterflies
Heinze, S., & Reppert, S. M. (2011) · Neuron · doi:10.1016/j.neuron.2010.12.025 · Cited by 268 · see citing works
The role of sensory adaptation in the retina
Laughlin, S. B. (1989) · Journal of Experimental Biology · doi:10.1242/jeb.146.1.39 · Cited by 256 · see citing works
Visual homing: an insect perspective
Zeil, J. (2012) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2011.12.008 · Cited by 248 · see citing works
Attractor Dynamics of Spatially Correlated Neural Activity in the Limbic System
Knierim, J. J., & Zhang, K. (2012) · Annual Review of Neuroscience · doi:10.1146/annurev-neuro-062111-150351 · Cited by 222 · see citing works
Object tracking in motion-blind flies
Bahl, A., Ammer, G., Schilling, T., & Borst, A. (2013) · Nature Neuroscience · doi:10.1038/nn.3386 · Cited by 152 · see citing works
Structure of the adult central complex in Drosophila: Organization of distinct neuronal subsets
Young, J. M., & Armstrong, J. D. (2009) · The Journal of Comparative Neurology · doi:10.1002/cne.22284 · Cited by 142 · see citing works
Animal Navigation: Path Integration, Visual Landmarks and Cognitive Maps
Collett, T. S., & Graham, P. (2004) · Current Biology · doi:10.1016/j.cub.2004.06.013 · Cited by 138 · see citing works
Functional dissection of circuitry in a neural integrator
Aksay, E., Olasagasti, I., Mensh, B. D., Baker, R., Goldman, M. S., & Tank, D. W. (2007) · Nature Neuroscience · doi:10.1038/nn1877 · Cited by 134 · see citing works
Collett, T. S. (2000) · Current Opinion in Neurobiology · doi:10.1016/s0959-4388(00)00150-1 · Cited by 131 · see citing works
Lin, C.-Y., Chuang, C.-C., Hua, T.-E., Chen, C., Dickson, B. J., Greenspan, R. J., & Chiang, A. (2013) · Cell Reports · doi:10.1016/j.celrep.2013.04.022 · Cited by 131 · see citing works
Real neuroscience in virtual worlds
Dombeck, D. A., & Reiser, M. B. (2011) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2011.10.015 · Cited by 115 · see citing works
Transformation of Polarized Light Information in the Central Complex of the Locust
Heinze, S., Gotthardt, S., & Homberg, U. (2009) · Journal of Neuroscience · doi:10.1523/jneurosci.1870-09.2009 · Cited by 113 · see citing works
Neural activity in the central complex of the cockroach brain is linked to turning behaviors
Guo, P., & Ritzmann, R. E. (2012) · Journal of Experimental Biology · doi:10.1242/jeb.080473 · Cited by 104 · see citing works
Yoshida, M., & Hasselmo, M. E. (2009) · The Journal of Neuroscience · doi:10.1523/jneurosci.5154-08.2009 · Cited by 94 · see citing works
Bausenwein, B., Müller, N. R., & Heisenberg, M. (1994) · The Journal of Comparative Neurology · doi:10.1002/cne.903400210 · Cited by 76 · see citing works
Persistence of orientation toward a temporarily invisible landmark in Drosophila melanogaster
Strauß, R., & Pichler, J. (1998) · Journal of Comparative Physiology A · doi:10.1007/s003590050190 · Cited by 75 · see citing works
Studying sensorimotor integration in insects
Huston, S. J., & Jayaraman, V. (2011) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2011.05.030 · Cited by 64 · see citing works
Self-citationEvolving a Neural Model of Insect Path Integration
Haferlach, T., Wessnitzer, J., Mangan, M., & Webb, B. H. (2007) · Adaptive Behavior · doi:10.1177/1059712307082080 · Cited by 53 · see citing works
Amplitude and dynamics of polarization-plane signaling in the central complex of the locust brain
Bockhorst, T., & Homberg, U. (2015) · Journal of Neurophysiology · doi:10.1152/jn.00742.2014 · Cited by 41 · see citing works
Kathman, N. D., Kesavan, M., & Ritzmann, R. E. (2014) · Journal of Experimental Biology · doi:10.1242/jeb.112391 · Cited by 28 · see citing works
A spiking network for spatial memory formation: Towards a fly-inspired ellipsoid body model
Arena, P., Maceo, S., Patané, L., & Strauß, R. (2013) · International Joint Conference on Neural Networks (IJCNN) · doi:10.1109/ijcnn.2013.6706882 · Cited by 18 · see citing works
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Spatial representation in the hippocampal formation: a history
Moser, E. I., Moser, M.-B., & McNaughton, B. L. (2017) · Nature Neuroscience · doi:10.1038/nn.4653 · Cited by 560 · see citing works
Toroidal topology of population activity in grid cells
Gardner, R. J., Hermansen, E., Pachitariu, M., Burak, Y., Baas, N. A., Dunn, B. A., Moser, M.-B., & Moser, E. I. (2022) · Nature · doi:10.1038/s41586-021-04268-7 · Cited by 428 · see citing works
Ring attractor dynamics in the <i>Drosophila</i> central brain
Kim, S. S., Rouault, H., Druckmann, S., & Jayaraman, V. (2017) · Science · doi:10.1126/science.aal4835 · Cited by 417 · see citing works
Self-citationWhat insects can tell us about the origins of consciousness.
Barron, A. B., & Klein, C. (2016) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1520084113 · Cited by 446 · see citing works
Whole-brain annotation and multi-connectome cell typing of Drosophila
Schlegel, P., Yin, Y., Bates, A. S., Dorkenwald, S., Eichler, K., Brooks, P., Han, D. S., Gkantia, M., dos Santos, M., Munnelly, E. J., et al. (2024) · Nature · doi:10.1038/s41586-024-07686-5 · Cited by 413 · see citing works
“However, the adult fly has richer behaviour, including complex motor control while walking or in flight 7 , courtship behaviour 8 , involved decision making 9 , flexible associative memory 10,11 , spatial learning 12 and complex 13,14 multisensory 15,16 navigation.”
BackgroundHulse, B. K., Haberkern, H., Franconville, R., Turner-Evans, D., Takemura, S.-Y., Wolff, T., Noorman, M., Dreher, M., Dan, C., Parekh, R., et al. (2021) · eLife · doi:10.7554/elife.66039 · Cited by 392 · see citing works
Self-citationA neural circuit architecture for angular integration in Drosophila
Green, J., Adachi, A., Shah, K. K., Hirokawa, J. D., Magani, P. S., & Maimon, G. (2017) · Nature · doi:10.1038/nature22343 · Cited by 331 · see citing works
An Anatomically Constrained Model for Path Integration in the Bee Brain
Stone, T., Webb, B., Adden, A., Weddig, N. B., Honkanen, A., Templin, R., Wcislo, W., Scimeca, L., Warrant, E., & Heinze, S. (2017) · Current Biology · doi:10.1016/j.cub.2017.08.052 · Cited by 377 · see citing works
Brain-wide mapping of neural activity controlling zebrafish exploratory locomotion
Dunn, T. W., Mu, Y., Narayan, S., Randlett, O., Naumann, E. A., Yang, C.-T., Schier, A. F., Freeman, J., Engert, F., & Ahrens, M. B. (2016) · eLife · doi:10.7554/elife.12741 · Cited by 312 · see citing works
Cellular diversity in the Drosophila midbrain revealed by single-cell transcriptomics
Croset, V., Treiber, C. D., & Waddell, S. (2018) · eLife · doi:10.7554/elife.34550 · Cited by 299 · see citing works
Angular velocity integration in a fly heading circuit
Turner-Evans, D., Wegener, S., Rouault, H., Franconville, R., Wolff, T., Seelig, J. D., Druckmann, S., & Jayaraman, V. (2017) · eLife · doi:10.7554/elife.23496 · Cited by 281 · see citing works
Self-citationFlyWire: online community for whole-brain connectomics
Dorkenwald, S., McKellar, C. E., Macrina, T., Kemnitz, N., Lee, K., Lu, R., Wu, J., Popovych, S., Mitchell, E., Nehoran, B., et al. (2022) · Nature Methods · doi:10.1038/s41592-021-01330-0 · Cited by 289 · see citing works
The insect central complex and the neural basis of navigational strategies
Honkanen, A., Adden, A., da Silva Freitas, J., & Heinze, S. (2019) · Journal of Experimental Biology · doi:10.1242/jeb.188854 · Cited by 238 · see citing works
Insects have the capacity for subjective experience
Klein, C., & Barron, A. B. (2016) · Animal Sentience · doi:10.51291/2377-7478.1113 · Cited by 243 · see citing works
Building an allocentric travelling direction signal via vector computation
Lyu, C., Abbott, L. F., & Maimon, G. (2022) · Nature · doi:10.1038/s41586-021-04067-0 · Cited by 215 · see citing works
Computational principles of memory
Chaudhuri, R., & Fiete, I. (2016) · Nature Neuroscience · doi:10.1038/nn.4237 · Cited by 213 · see citing works
Principles of insect path integration
Heinze, S., Narendra, A., & Cheung, A. (2018) · Current Biology · doi:10.1016/j.cub.2018.04.058 · Cited by 195 · see citing works
Generation of stable heading representations in diverse visual scenes
Kim, S. S., Hermundstad, A. M., Romani, S., Abbott, L. F., & Jayaraman, V. (2019) · Nature · doi:10.1038/s41586-019-1767-1 · Cited by 172 · see citing works
Self-citationSensorimotor experience remaps visual input to a heading-direction network
Fisher, Y. E., Lu, J., D’Alessandro, I., & Wilson, R. I. (2019) · Nature · doi:10.1038/s41586-019-1772-4 · Cited by 165 · see citing works
Wolff, T., & Rubin, G. M. (2018) · Journal of Comparative Neurology · doi:10.1002/cne.24512 · Cited by 156 · see citing works
Sun Navigation Requires Compass Neurons in Drosophila
Giraldo, Y. M., Leitch, K. J., Ros, I. G., Warren, T. L., Weir, P. T., & Dickinson, M. H. (2018) · Current Biology · doi:10.1016/j.cub.2018.07.002 · Cited by 161 · see citing works
The Neuroanatomical Ultrastructure and Function of a Biological Ring Attractor
Turner-Evans, D. B., Jensen, K. T., Ali, S., Paterson, T., Sheridan, A., Ray, R. P., Wolff, T., Lauritzen, J. S., Rubin, G. M., Bock, D. D., et al. (2020) · Neuron · doi:10.1016/j.neuron.2020.08.006 · Cited by 168 · see citing works
Self-citationThe neuroanatomical ultrastructure and function of a biological ring attractor
Turner-Evans, D. B., Jensen, K. T., Ali, S., Paterson, T., Sheridan, A., Ray, R. P., Wolff, T., Lauritzen, S., Rubin, G. M., Bock, D., et al. (2019) · doi:10.1101/847152 · Cited by 166 · see citing works
Self-citationTransforming representations of movement from body- to world-centric space
Lu, J., Behbahani, A. H., Hamburg, L., Westeinde, E. A., Dawson, P. M., Lyu, C., Maimon, G., Dickinson, M. H., Druckmann, S., & Wilson, R. I. (2022) · Nature · doi:10.1038/s41586-021-04191-x · Cited by 152 · see citing works
Mechanisms Underlying the Neural Computation of Head Direction
Hulse, B. K., & Jayaraman, V. (2020) · Annual Review of Neuroscience · doi:10.1146/annurev-neuro-072116-031516 · Cited by 130 · see citing works
Self-citationIdiothetic Path Integration in the Fruit Fly Drosophila melanogaster
Kim, I. S., & Dickinson, M. H. (2017) · Current Biology · doi:10.1016/j.cub.2017.06.026 · Cited by 119 · see citing works
Sexual arousal gates visual processing during Drosophila courtship
Hindmarsh Sten, T., Li, R., Otopalik, A., & Ruta, V. (2021) · Nature · doi:10.1038/s41586-021-03714-w · Cited by 140 · see citing works
A neural heading estimate is compared with an internal goal to guide oriented navigation
Green, J., Vijayan, V., Mussells Pires, P., Adachi, A., & Maimon, G. (2019) · Nature Neuroscience · doi:10.1038/s41593-019-0444-x · Cited by 137 · see citing works
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Okubo, T. S., Patella, P., D’Alessandro, I., & Wilson, R. I. (2020) · Neuron · doi:10.1016/j.neuron.2020.06.022 · Cited by 114 · see citing works
Network statistics of the whole-brain connectome of Drosophila
Lin, A., Yang, R., Dorkenwald, S., Matsliah, A., Sterling, A. R., Schlegel, P., Yu, S.-C., McKellar, C. E., Costa, M., Eichler, K., et al. (2024) · Nature · doi:10.1038/s41586-024-07968-y · Cited by 147 · see citing works
The geometry of decision-making in individuals and collectives
Sridhar, V. H., Li, L., Gorbonos, D., Nagy, M., Schell, B. R., Sorochkin, T., Gov, N. S., & Couzin, I. D. (2021) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2102157118 · Cited by 120 · see citing works
Optimal multiguidance integration in insect navigation
Hoinville, T., & Wehner, R. (2018) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1721668115 · Cited by 116 · see citing works
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Bidaye, S. S., Laturney, M., Chang, A. K., Liu, Y., Bockemühl, T., Büschges, A., & Scott, K. (2020) · Neuron · doi:10.1016/j.neuron.2020.07.032 · Cited by 109 · see citing works
A visual pathway for skylight polarization processing in Drosophila
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Schlegel, P., Yin, Y., Bates, A. S., Dorkenwald, S., Eichler, K., Brooks, P., Han, D. S., Gkantia, M., dos Santos, M., Munnelly, E. J., et al. (2023) · doi:10.1101/2023.06.27.546055 · Cited by 102 · see citing works
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Kakaria, K. S., & de Bivort, B. L. (2017) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2017.00008 · Cited by 89 · 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
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
The Central Complex as a Potential Substrate for Vector Based Navigation
Le Moël, F., Stone, T., Lihoreau, M., Wystrach, A., & Webb, B. (2019) · Frontiers in Psychology · doi:10.3389/fpsyg.2019.00690 · Cited by 86 · see citing works
Vinepinsky, E., Cohen, L., Perchik, S., Ben-Shahar, O., Donchin, O., & Segev, R. (2020) · Scientific Reports · doi:10.1038/s41598-020-71217-1 · Cited by 87 · see citing works
Comparative Neuroanatomy of the Lateral Accessory Lobe in the Insect Brain
Namiki, S., & Kanzaki, R. (2016) · Frontiers in Physiology · doi:10.3389/fphys.2016.00244 · Cited by 78 · 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
Flexible 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
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
A Snapshot-Based Mechanism for Celestial Orientation
el Jundi, B., Foster, J. J., Khaldy, L., Byrne, M. J., Dacke, M., & Baird, E. (2016) · Current Biology · doi:10.1016/j.cub.2016.03.030 · Cited by 76 · 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
The Drosophila Split Gal4 System for Neural Circuit Mapping
Luan, H., Diao, F., Scott, R. L., & White, B. H. (2020) · Frontiers in Neural Circuits · doi:10.3389/fncir.2020.603397 · Cited by 73 · 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
A computational model of the integration of landmarks and motion in the insect central complex
Cope, A. J., Sabo, C., Vasilaki, E., Barron, A. B., & Marshall, J. A. R. (2017) · PLOS ONE · doi:10.1371/journal.pone.0172325 · Cited by 51 · 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
A unified mechanism for innate and learned visual landmark guidance in the insect central complex
Goulard, R., Buehlmann, C., Niven, J. E., Graham, P., & Webb, B. (2021) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1009383 · Cited by 51 · see citing works
A model of egocentric to allocentric understanding in mammalian brains
Uria, B., Ibarz, B., Banino, A., Zambaldi, V., Kumaran, D., Hassabis, D., Barry, C., & Blundell, C. (2020) · doi:10.1101/2020.11.11.378141 · Cited by 48 · see citing works
How foresight might support the behavioral flexibility of arthropods
Perry, C. J., & Chittka, L. (2019) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2018.10.014 · Cited by 54 · see citing works
Hulse, 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
“We began by assuming that the EPG neurons have an activity bump, similar to that observed experimentally (Seelig and Jayaraman, 2015).”
BackgroundResultInfluentialSelf-citationFine-grained descending control of steering in walking Drosophila
Yang, H. H., Brezovec, B. E., Serratosa Capdevila, L., Vanderbeck, Q. X., Adachi, A., Mann, R. S., & Wilson, R. I. (2024) · Cell · doi:10.1016/j.cell.2024.08.033 · Cited by 55 · see citing works
Neural networks for navigation: From connections to computations
Wilson, R. I. (2023) · Annual Review of Neuroscience · doi:10.1146/annurev-neuro-110920-032645 · Cited by 38 · see citing works
A neural circuit architecture for rapid learning in goal-directed navigation
Dan, C., Hulse, B. K., Kappagantula, R., Jayaraman, V., & Hermundstad, A. M. (2024) · Neuron · doi:10.1016/j.neuron.2024.04.036 · Cited by 51 · see citing works
“36,44,48 Modeling Determining the optimal policy for maintaining a goal heading Figure 3D shows the drift rate and average duration of fixations, and the turn bias of saccades, that result from training a flexible RL agent to exhibit a preference for a goal heading.”
BackgroundMethodInfluentialOn the role of the head ganglia in posture and walking in insects
Emanuel, S., Kaiser, M., Pflueger, H.-J., & Libersat, F. (2020) · Frontiers in Physiology · doi:10.3389/fphys.2020.00135 · Cited by 31 · see citing works
Bayesian inference in ring attractor networks
Kutschireiter, A., Basnak, M. A., Wilson, R. I., & Drugowitsch, J. (2023) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2210622120 · Cited by 36 · see citing works
Neural circuit mechanisms for steering control in walking Drosophila
Rayshubskiy, A., Holtz, S. L., Bates, A. S., Vanderbeck, Q. X., Serratosa Capdevila, L., Rockwell, V., & Wilson, R. (2025) · eLife · doi:10.7554/elife.102230 · Cited by 27 · see citing works
An Analysis of a Ring Attractor Model for Cue Integration
Sun, X., Mangan, M., & Yue, S. (2018) · Lecture Notes in Computer Science · doi:10.1007/978-3-319-95972-6_49 · Cited by 27 · see citing works
Lazar, A. A., Liu, T., Turkcan, M. K., & Zhou, Y. (2021) · eLife · doi:10.7554/elife.62362 · Cited by 21 · see citing works
Towards a common terminology for arthropod spatial orientation
Grob, R., el Jundi, B., & Fleischmann, P. N. (2021) · Ethology Ecology & Evolution · doi:10.1080/03949370.2021.1905075 · Cited by 22 · see citing works
Batchelor, A. V., & Wilson, R. I. (2019) · Journal of Experimental Biology · doi:10.1242/jeb.191213 · Cited by 20 · see citing works
Nakayama, K., Moher, J., & Song, J.-H. (2023) · Annual Review of Psychology · doi:10.1146/annurev-psych-021422-043229 · Cited by 21 · see citing works
Maintaining a stable head direction representation in naturalistic visual environments
Haberkern, H., Chitnis, S. S., Hubbard, P. M., Goulet, T., Hermundstad, A. M., & Jayaraman, V. (2022) · doi:10.1101/2022.05.17.492284 · Cited by 21 · see citing works
Self-citationA neural circuit architecture for rapid behavioral flexibility in goal-directed navigation
Dan, C., Hulse, B. K., Kappagantula, R., Jayaraman, V., & Hermundstad, A. M. (2021) · doi:10.1101/2021.08.18.456004 · Cited by 51 · see citing works
“This triggers a competition between two sets of compass neurons that are activated by the same ring neurons [16, 31, 34, 71], and the bump jumps across the EB to the location that is most weakly inhibited by the same active ring neurons (Fig 2d, right column).”
BackgroundMethodResultInfluentialSelf-citationFine-grained descending control of steering in walking <i>Drosophila</i>
Yang, H. H., Brezovec, L. E., Serratosa Capdevila, L., Vanderbeck, Q. X., Adachi, A., Mann, R. S., & Wilson, R. I. (2023) · doi:10.1101/2023.10.15.562426 · Cited by 55 · see citing works
A model of cue integration as vector summation in the insect brain
Mitchell, R., Shaverdian, S., Dacke, M., & Webb, B. (2023) · Proceedings of the Royal Society B: Biological Sciences · doi:10.1098/rspb.2023.0767 · Cited by 13 · see citing works
“In order to account for the results of [14] (E-PG bump retention in darkness and E-PG visual capture) as well as our own proposed learning routine (where E-PGs are driven primarily by angular velocity input), input to the E-PGs is context dependent.”
BackgroundConverting an allocentric goal into an egocentric steering signal
Pires, P. M., Abbott, L. F., & Maimon, G. (2022) · doi:10.1101/2022.11.10.516026 · Cited by 110 · see citing works
Multimodal cue integration and learning in a neural representation of head direction
Basnak, M. A., Kutschireiter, A., Okubo, T. S., Chen, A., Gorelik, P., Drugowitsch, J., & Wilson, R. I. (2025) · Nature Neuroscience · doi:10.1038/s41593-024-01823-z · Cited by 13 · see citing works
“All the HD cells in this network can be imaged simultaneously as a head-fixed fly navigates in a virtual reality environment 33,34 .”
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. (2022) · doi:10.1101/2022.11.10.516039 · Cited by 85 · see citing works
Weighted cue integration for straight-line orientation
Shaverdian, S., Dirlik, E., Mitchell, R., Tocco, C., Webb, B., & Dacke, M. (2022) · iScience · doi:10.1016/j.isci.2022.105207 · Cited by 9 · see citing works
Phenomenal interface theory: a model for basal consciousness
Klein, C., & Barron, A. B. (2025) · Philosophical Transactions of the Royal Society B: Biological Sciences · doi:10.1098/rstb.2024.0301 · Cited by 8 · see citing works
“Processing within the central complex enables an insect to establish and hold a heading relative to an external navigational reference [45,51].”
Neural circuit mechanisms for steering control in walking Drosophila
Rayshubskiy, A., Holtz, S. L., Bates, A., Vanderbeck, Q. X., Capdevila, L. S., & Wilson, R. I. (2024) · doi:10.7554/elife.102230.1 · Cited by 5 · see citing works
- Authors
- Seelig, J. D., & Jayaraman, V.
- Affiliations
- Howard Hughes Medical Institute (US)
- Corresponding author
- Jayaraman, Vivek
- Venue
- Nature
- Published
- 2015-05
- Volume
- 521
- Issue
- 7551
- Pages
- 186-191
- ISSN
- 0028-0836 (ISSN-L)
- 1476-4687
- PMC
- PMC4704792
- OpenAlex
- W1503270516
- Cited by
- 718
- References
- 47
- Citations by year
- 2026: 60 · 2025: 77 · 2024: 73 · 2023: 63 · 2022: 69 · 2021: 71 · 2020: 61 · 2019: 70 · 2018: 59 · 2017: 66 · 2016: 37 · 2015: 12
- FWCI
- 27.08
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- 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)
- MPG.PuRe (Max Planck Society) (submittedVersion)
- Funders
- Howard Hughes Medical Institute
- Topics
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurobiology and Insect Physiology Research
- Physical Sciences › Engineering › Automotive Engineering › Spatial Cognition and Navigation
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Keywords
- head direction cells, ellipsoid body, Drosophila melanogaster, spatial navigation, persistent activity, ring attractor, self-motion cues, insect brain, population coding
- MeSH terms
- Animals, Brain, Calcium, Cues, Darkness, Dendrites, Drosophila melanogaster, Female, Head, Neurons, Orientation, Photic Stimulation, Rotation, Space Perception, Walking, Spatial Navigation
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