FlyBase: A Database of Drosophila Genes & Genomes
The official fly gene and genome database, for looking up genes, strains, life history, and literature โ a foundational tool for learning fly biology.
The fruit fly Drosophila melanogaster is one of the most important model organisms in neuroscience and genetics: it is small, has a short life cycle, has mature genetic tools, and now has a fully sequenced whole-brain connectome. The resources below take you from the fly's basic biology and brain structure to the latest connectome data, introducing this 'condensed brain.'
The official fly gene and genome database, for looking up genes, strains, life history, and literature โ a foundational tool for learning fly biology.
An HHMI BioInteractive animation explaining how clock genes such as period and timeless use negative feedback loops to control the fly's day/night rhythm, closely related to this project's 'rhythm' theme.
The flagship 2024 Nature paper by Dorkenwald et al. presenting the whole-brain connectome of an adult female fruit fly, with 139,255 neurons and over 50 million chemical synapses โ the authoritative data source for the scale of the fly brain.
An interactive fly neuroanatomy atlas integrating over 610,000 3D neural images, connectome data, and gene-expression data; you can browse brain regions and neuron morphology online, making it a good entry point for an overview of brain structure.
An open-access 2025 graphical review in Current Research in Insect Science that lays out the structure, circuitry and associated behaviours of the adult fly brain for newcomers, with pointers to further resources for each topic. A fast way to build a whole-brain overview.
A 2025 Science Advances paper that used the connectome to find SLP316, the main postsynaptic partner of the fly's sLNv pacemaker neurons. Silencing or exciting SLP316 shortens or lengthens the free-running period, showing that synaptic wiring feeds back on the clock's own parameters. Directly relevant to this project's rhythm model.
The official Nobel press release explaining how Hall, Rosbash and Young used the fruit fly to uncover the negative feedback loop of period, timeless and related genes that underlies the molecular circadian clock. A first-hand, authoritative account of the rhythm mechanism.
A 2026 Cell paper reporting MaleCNS, the first reconstruction of the entire Drosophila male central nervous system as a single connectome โ brain and ventral nerve cord together, 166,700 neurons โ a milestone that extends the "brain map" into a full-body nervous-system map.
Since 2018, Google Research has worked with Janelia/HHMI, the University of Cambridge and Princeton to build fly connectomes step by step: from the hemibrain to the female whole brain (FlyWire/FAFB), then the female brain-and-nerve-cord BANC connectome released in June 2026, and finally the complete male central nervous system (MaleCNS) released in September 2026. The resources below cover official blog posts, Nature and Cell papers, online browsers and simulation platforms, and form the technical and data base for this digital-twin project.
Google's announcement of 3 September 2026: with HHMI Janelia, the University of Cambridge and the MRC LMB, they completed the full connectome of the male fly brain plus ventral nerve cord, over 166,000 neurons and about 11,700 cell types, with the paper published the same day in Cell; the data are CC-BY at male-cns.janelia.org (neuPrint, Clio, Cell Type Explorer); version of record: Berg et al., Cell 189(18):5504โ5526, 3 Sept 2026, doi:10.1016/j.cell.2026.08.015 (166,700 neurons, 11,710 cell types).
A 2019 Google Research blog post describing how Flood-Filling Networks (a machine-learning algorithm) were used to automatically reconstruct roughly 100,000 neurons of the fly whole brain (the FAFB dataset), and introducing the Neuroglancer online viewer.
A 2020 blog post from Google Research and the Janelia FlyEM team releasing the hemibrain connectome, covering about 25,000 neurons and over 20 million synaptic connections, including the mushroom body and central complex. The word 'largest' in the title reflects 2020; the FlyWire whole brain, BANC and MaleCNS connectomes have since surpassed it.
A September 2026 Google Research blog post on the complete male Drosophila central nervous system connectome, produced with the Janelia FlyEM and Cambridge connectomics teams. It spans brain, optic lobes and ventral nerve cord with about 166,000 neurons and over a hundred million synaptic connections, the largest complete CNS map by neuron count so far (the female BANC has roughly 160,000 neurons (155,916 proofread)).
The 2024 Nature flagship paper by Dorkenwald et al. releasing the FlyWire adult female fruit fly whole-brain connectome (139,255 neurons, over 50 million synapses), with cell-type and neurotransmitter-prediction annotations.
A 2024 Nature paper by Schlegel et al. establishing a systematic, hierarchical cell-type annotation for the FlyWire whole-brain connectome, labeling 8,453 cell types โ an important reference for understanding neuron classification across brain regions.
The official FlyWire connectome browsing platform Codex (developed by the Princeton Neuroscience Institute), letting you search neurons with free-text or structured queries, filter by cell type (e.g. Kenyon cells, KC), and inspect neural connections and synaptic paths in 3D without downloading large datasets.
Janelia's official connectome query tool neuPrint, for looking up connection partners, connection strength, and morphology of specific neurons in datasets like the hemibrain, with both a web interface and programmatic Python access.
The official website for the male fruit fly central nervous system connectome (v1.0 released in 2026), built by Janelia FlyEM, the MRC Laboratory of Molecular Biology at Cambridge, and Google Research, offering interactive tools such as a Cell Type Explorer and Dimorphism Explorer, sex-difference comparisons, and data downloads under a CC-BY license.
A fly digital-twin simulation platform developed by EPFL's Neuroengineering Lab, combining a biomechanical exoskeleton built from micro-CT scans, vision and olfaction sensing modules, and a hierarchical neural controller โ an important reference example for this rhythm-decision digital-twin bio-inspired platform project.
The flagship MaleCNS paper, published in Cell on 3 September 2026. It fully reconstructs and proofreads 166,700 neurons and 11,710 cell types across the male brain and nerve cord, annotates fruitless/doublesex expression, and makes the first synapse-resolution male-versus-female comparison: 8,069 isomorphic, 138 dimorphic, 289 male-specific and 71 female-specific types.
The BANC connectome paper, published in Nature on 8 June 2026. It is the first reconstruction to join the brain and ventral nerve cord of an adult female fly in a single electron-microscopy volume, roughly 160,000 neurons (155,916 proofread). Effector neurons are mostly driven by sensory neurons in the same body part, forming local loops that ascending and descending neurons link into behaviour-centric modules: a distributed, parallel and embodied architecture.
FlyWire's official BANC portal: how to browse the female brain-and-nerve-cord connectome in Codex, plus the application route for proofreading and annotation volunteers and the citation guidelines.
The FlyWire blog announcement of 3 November 2025: with collaborators at Harvard Medical School they released BANC, a synapse-resolution wiring diagram covering the entire central nervous system of an adult fly, free to use and open for the community to keep improving in Codex.
Virtual Fly Brain's official news feed, listing data and tool updates from 2024 to 2026 in date order: NBLAST similarity extended to the MaleCNS optic lobe and FlyWire datasets in 2025, BANC tour videos and MaleCNS connectome videos in 2026, and a free self-led online workshop.
The fly visual system is built around the optic lobe, with layered neuropils (medulla, lobula, lobula plate) where motion detectors such as T4/T5 and looming detectors such as LC4/LPLC2 support motion vision and escape. FlyWire mapped the roughly 40,000 optic-lobe neurons of the female fly; in 2025 Janelia published a complete inventory of one male optic lobe (about 53,000 neurons in 732 cell types), and the 2026 MaleCNS connectome traces visual signals all the way into the central brain, so function can be predicted directly from structure.
The FlyWire Codex provides a complete connectomic database of fly optic-lobe neurons, letting you search and browse the cell types and wiring information of over 40,000 visual neurons.
A 2024 Nature FlyWire connectome paper that, by analyzing the wiring structure of Dm3 and TmY cells, successfully predicted their function in shape vision and direction detection.
A classic Nature paper finding that LPLC2 neurons respond with high selectivity to an approaching object via a four-direction radial motion opponency mechanism, triggering escape behavior.
An eLife paper exploring how looming-detection neurons such as LC4 convert visual looming information into a spatial position signal in the central brain, guiding directional escape responses.
A PNAS paper showing that fly motion detection relies entirely on brightness contrast from R1-R6 photoreceptors, operating fully independently of the R7/R8 photoreceptors responsible for color vision.
An official Google Research blog post introducing the complete male fly central nervous system connectome, completed with HHMI Janelia, covering the optic lobe along with a total of 166,000 neurons.
One of the three MaleCNS companion papers (Cell, 3 September 2026). It traces signals from photoreceptors through the optic lobes, which hold two thirds of the brain's neurons, and onward into the central brain, revealing a multilayered set of pathway classes. It also predicts receptive-field structure and feature biases for thousands of uncharacterised neuron types, consistent with existing physiology.
Nern and colleagues, Nature 2025: a FIB-SEM connectome of the right optic lobe of a male fly, classifying roughly 53,000 neurons into 732 cell types, about half of them newly named, and released alongside matched split-GAL4 driver lines. The authoritative catalogue for visual cell types.
A 2025 Nature paper that reconstructed hundreds of T4 neurons and found that the geometry of the compound eye sets the systematic variation in preferred directions of direction-selective neurons across the whole visual field. The shape of the eye directly shapes how optic flow is decoded.
The fruit fly has no ears; instead, Johnston's organ in the second segment of the antenna serves as its receptor for hearing and mechanosensation, detecting vibration of the antennal arista to sense courtship song, wind direction, and gravity. The signal is relayed through the antennal mechanosensory and motor center (AMMC) to higher brain regions, forming a dedicated auditory-processing circuit.
A Nature paper showing how different subpopulations of neurons within Johnston's organ separately handle gravity sensing and hearing, with central projections resembling the vestibular and auditory pathways of mammals.
An eLife paper identifying the male-specific aPN1-vPN1-pC1 courtship-song processing circuit, explaining how the auditory signal is translated and triggers courtship behavior.
A Virtual Fly Brain database page detailing the anatomy of Johnston's organ, the arrangement of its roughly 477 sensory neurons, and their functional grouping for detecting sound, wind direction, and gravity.
A Virtual Fly Brain database page for an AMMC neuron, providing the anatomical morphology and connectome data of this auditory relay neuron, with a link to a 3D viewer.
A 2025 iScience paper showing that auditory sensory neurons in the female fly's Johnston's organ carry abundant dopamine receptors, and that Dop1R2 boosts song responses only in unmated females. Mating state therefore tunes hearing right at the first sensory stage, via dopamine.
Fly olfactory receptor neurons (ORNs) are distributed across the antennae and maxillary palps and relay odor signals into roughly 50 glomeruli in the antennal lobe; projection neurons (PNs) then send this information to the mushroom body and lateral horn for learned and innate processing. A team led by Professor Ann-Shyn Chiang at National Tsing Hua University produced the world's first complete fly olfactory neural map.
An eLife paper using FIB-SEM to fully reconstruct all neurons and synaptic connections of a single glomerulus, VA1v, including olfactory receptor neurons, projection neurons, and local interneurons.
An eLife paper analyzing the complete olfactory system in the hemibrain connectome, classifying neurons and finding a high degree of stereotypy โ structural consistency โ in olfactory circuits across individuals.
An eLife paper explaining how the anatomical arrangement of antennal-lobe projection neurons encodes odor responses and relays that information to the lateral horn for innate behavioral decisions.
A Virtual Fly Brain database page showing the location of the DM2 glomerulus in the antennal lobe and the anatomical details of its Or22-receptor ORN input, with a 3D visualization tool.
A 2026 Nature paper using a virtual-reality odour landscape to show that plume tracking is not purely reflexive. Flies edge-track along the plume boundary, using vector computations in the central complex, with FC2 neurons storing and continuously updating the direction back to the edge. It links olfaction to navigational memory.
Gustatory receptor neurons (GRNs) sit mainly on the labellum of the proboscis, detecting sugar, bitter, salt and water, and project to the subesophageal zone (SEZ) of the brain. Connectome work shows the different taste modalities stay largely separate in the SEZ, forming parallel circuits; the complete gustatory connectome published in 2026 traces whole sensory-to-motor paths onward into feeding, foraging and courtship circuits.
An eLife paper using electron microscopy to reconstruct the connectome of 144 taste neurons, finding that 79% of synaptic connections occur within the same taste modality, showing that sugar, bitter, salt, and water signals each form independent circuits in the SEZ.
A Scientific Reports paper using whole-brain connectome data to trace how taste signals travel from sensory neurons through the SEZ to higher brain regions, comparing the projection differences between second- and third-order taste neurons.
An eLife paper identifying three taste projection neurons (TPN1-3), of which TPN3 carries bitter-taste information and projects to the mushroom body learning center, and is required for forming conditioned taste-aversion learning.
A Virtual Fly Brain database page describing the anatomy of labellar taste-peg chemosensory neurons and their connection data projecting to the subesophageal zone, with associated FlyWire connectome imagery.
A Virtual Fly Brain page introducing the sensory neuron in the labellar taste bristles that detects high salt concentration, describing its expression of the ppk23 receptor and distribution in the labellum's taste bristles.
One of the three MaleCNS companion papers (Cell, 3 September 2026). It is the first complete reconstruction of gustatory receptor neurons in a single electron-microscopy volume spanning brain, cervical connective and ventral nerve cord, integrated into a cross-sex pan-CNS gustatory connectome. It traces full paths from taste input to feeding motor neurons across four circuit types: nutrient assessment, motor control, neuroendocrine regulation and courtship.
Fly walking, flight, and escape movements are controlled by motor neurons and central pattern generator (CPG) circuits in the thoracic ganglia (ventral nerve cord, VNC). Recent electron-microscopy connectome reconstructions and neuromechanical simulation platforms have, for the first time, let scientists study at single-synapse resolution how the six-legged tripod gait and flight motor circuits are generated.
A paper from an EPFL team introducing the second-generation neuromechanical fly simulation platform, integrating vision, olfaction, and motor feedback to reproduce walking, obstacle avoidance, and odor-guided navigation.
The official project site of EPFL's Ramdya Lab, offering the browser-interactive fly biomechanics simulator FlyGym for observing gait and terrain-adaptation behavior.
The official NeuroMechFly source-code repository, including the biomechanical skeleton, muscle models, and neural controller, allowing gait to be reproduced and optimized.
A 2024 Nature paper using electron microscopy to reconstruct the female fly ventral nerve cord (VNC), mapping the muscle correspondence of leg and wing motor neurons and identifying the circuit that coordinates takeoff.
An analysis paper on the complete male ventral nerve cord connectome (MANC), showing how descending neurons drive motor neurons through intermediary layers rather than direct connections, delineating functional modules for walking, flight, and more.
Janelia's official connectome query platform, letting you interactively search neuron connections in the MANC ventral nerve cord and hemibrain datasets โ well suited for looking up locomotion-related circuits.
The MANC connectome paper from the Janelia FlyEM team, eLife 2024. It densely reconstructs the entire male ventral nerve cord, the fly's equivalent of a spinal cord, including the motor neurons that drive the legs and wings. It is the dataset that lets brain signals be traced all the way to muscle, and the foundation the later whole-CNS MaleCNS connectome was built on.
A 2026 Nature Communications paper from the Tuthill lab showing that a hair-plate proprioceptor on the fly's leg coxa detects when the leg nears the limit of forward movement, and using electron-microscopy reconstruction to show its axons excite posterior-leg motor neurons while inhibiting anterior ones, preventing overextension.
The mushroom body is the fly's center for associative learning and memory, made up of about 2,000 Kenyon cells, paired with roughly twenty types of dopaminergic neurons that convey reward or punishment signals, forming plastic synapses across fifteen lobe compartments so the fly can link odors with experience.
A classic 2014 eLife paper fully mapping the connectivity of the mushroom body's roughly 2,200 neurons, including 7 types of Kenyon cells, 21 types of output neurons, and 20 types of dopaminergic neurons, revealing the circuit logic of associative learning.
This study found reciprocal synapses between Kenyon cells and dopaminergic neurons, where cholinergic input from Kenyon cells amplifies dopamine release, forming the positive feedback loop required for learning.
The Virtual Fly Brain entry for the adult fly Kenyon cell, integrating connectome, gene-expression, and 3D neuroanatomical data for interactively browsing mushroom-body cell morphology.
A 2026 Nature Neuroscience paper showing that forgotten aversive memories persist as silent traces in behaviourally neutral mushroom body output neurons. Reminder cues reinstate the original trace and recover the memory, but misleading reminders create false memories instead, with the two outcomes gated by separable dopaminergic pathways.
The ventral nerve cord (VNC) is the fly's "spinal cord": roughly 15,000-23,000 neurons that integrate descending commands and sensory feedback to drive whole-body actions such as six-legged walking, flight, grooming, courtship, and egg-laying. Between 2024 and 2026, connectome datasets such as MANC, BANC, and MaleCNS reconstructed the entire thoracic-abdominal ganglion at synaptic resolution for the first time, letting scientists trace signals all the way from brain to muscle.
Janelia FlyEM's densely reconstructed connectome of the male Drosophila ventral nerve cord (MANC), covering roughly 23,000 neurons and about 10 million synapses, including the motor neurons that drive the legs and wings โ the key dataset for tracing descending signals from brain to muscle (eLife reviewed preprint, first version 2024).
Janelia FlyEM's official MANC project page, describing the June 2023 release built with Cambridge and Google Research: a dense reconstruction of about 23,000 neurons, 10 million pre-synaptic sites, and 74 million post-synaptic densities in the male VNC.
A deep link into Janelia's neuPrint explorer preset to the MANC v1.0 dataset, for directly searching ventral-nerve-cord motor neurons, descending neurons, and their synaptic partners โ useful for looking up leg and wing circuits.
A 2020 Neuron consensus paper from multiple labs extending the adult-brain nomenclature of Ito et al. (2014) to the thoracic and abdominal ganglia, defining neuropil names such as NTct, WTct, and LegNp T1-T3 โ the naming standard behind this platform's VNC neuropil list.
A 2026 Nature paper from the FlyWire consortium reporting the BANC (brain-and-cord) connectome, the first single reconstruction spanning both the female fly's brain and ventral nerve cord โ roughly 10^8 synaptic connections โ used to analyze how descending, ascending, and motor neurons distribute control of whole-body behavior.
A 2026 Cell paper presenting MaleCNS, the connectome of the entire Drosophila male central nervous system โ brain and nerve cord together, 166,700 fully proofread neurons across 11,710 cell types โ and the first comprehensive male-versus-female brain connectome comparison at synaptic resolution.
A 2025 Nature paper integrating three separate electron-microscopy datasets to systematically compare descending neurons (DNs), ascending neurons (ANs), and sensory ascending neurons crossing the neck connective โ the anatomical and information bottleneck between brain and ventral nerve cord (roughly 1,300 descending-neuron types).
A 2024 Nature paper reconstructing the female Drosophila VNC connectome โ about 45 million synapses and 14,600 neurons โ and using genetic labeling to map leg and wing motor neurons to the individual muscles each one drives, the first complete neuron-to-muscle map.
An analysis of the MANC connectome showing how descending neurons drive motor output: most descending input reaches motor neurons indirectly, through an intermediate layer of premotor interneuron communities organized into functional modules for walking, flight, and other behaviors.
A landmark 2018 Neuron paper from the Tuthill lab using in vivo calcium imaging of the femoral chordotonal organ to show how different proprioceptor classes encode leg joint angle, direction, and vibration โ the functional foundation later connectome studies of VNC proprioceptive circuits build on.
A 2026 Current Biology paper building a neural connectivity atlas for fly flight control, showing how topographically organized sensory maps correspond to the motor output of wing-steering muscles โ the circuit logic behind sub-millisecond flight stability adjustments.
A 2026 eLife paper from the Seeds lab reconstructing the complete mechanosensory connectome of the Drosophila head, showing how touch-sensing neurons are organized somatotopically into circuits that let the fly aim grooming precisely at the stimulated body location.
A 2020 Nature paper identifying the oviposition descending neurons (oviDNs) that are necessary and sufficient to trigger egg laying in Drosophila, and showing how post-mating sex-peptide signaling releases the inhibition that normally keeps oviDNs silent in virgin females.
A 2024 Nature paper building a leaky integrate-and-fire computational model of the entire fly brain connectome, which accurately predicts how activating sugar- or water-sensing gustatory neurons drives the motor output for proboscis extension and grooming.
Virtual Fly Brain's current standard adult ventral-nerve-cord template (JRCVNC2018 Unisex), the VNC counterpart to the JRC2018 brain template, built from combined male and female confocal images and used to align VNC connectome datasets to a common coordinate space.
The second-generation whole-body biomechanical simulation platform from EPFL, integrating vision, olfaction, and motor feedback to reproduce walking, obstacle avoidance, and odor-guided navigation โ a full-body digital-twin tool for testing hypotheses about VNC motor circuits.
Courtship, Aggression & Sex Differences
The fruitless (fru) and doublesex (dsx) genes determine the sex differences in fly neural circuits, and P1 neurons integrate multiple sensory signals to drive male courtship behavior. The male and female whole-brain connectomes published between 2024 and 2026 let scientists, for the first time, see the neural basis of courtship, aggression, and sexual dimorphism at synaptic resolution.
Sexual dimorphism in the complete connectome of the Drosophila male central nervous system
The MaleCNS team's October 2025 bioRxiv preprint comparing sexual dimorphism across the male and female central nervous system connectomes; the final version appeared in Cell on 3 September 2026.
Researchers reveal connectome of the male fruit fly central nervous system
A Janelia news release (October 2025) announcing the complete male fly central nervous system connectome, which for the first time lets scientists compare male and female brain circuitry at scale to study courtship and aggression.
Why do flies fall in love? Researchers tease out the signals behind fruit fly courtship songs
A Janelia news piece introducing the neural circuit behind male courtship song, including actual video and audio recordings of male courtship behavior and song.
A connectomics milestone: Mapping the complete male fruit fly brain
An official Google Research blog post describing the completion, with HHMI Janelia, of the whole male fly brain connectome, containing over 166,000 neurons and 125 million synaptic connections.
Drosulfakinin signaling in fruitless circuitry antagonizes P1 neurons to regulate sexual arousal in Drosophila
A Nature Communications paper showing that four pairs of fruitless neurons expressing the neuropeptide drosulfakinin antagonize P1 neuron activity, regulating the level of sexual arousal in male flies.
Male CNS Connectome Explorer
The official portal for the male fly central nervous system connectome, offering neuPrint queries, the Clio cell-type annotation tool, and a dimorphism explorer for comparing male/female brain differences.
Networks of sexually dimorphic neurons that regulate social behaviors in Drosophila
One of the three MaleCNS companion papers (Current Biology, 3 September 2026). Using the connectome it splits the male pC1/pC1x population into 48 cell types embedded in an expansive, highly intertwined male-specific network, and it delivers genetic driver lines showing that different pC1 types control social acoustic signalling and male-male interaction.
Two classes of amine/glutamate multi-transmitter neurons innervate Drosophila internal male reproductive organs
A 2026 eLife paper identifying two classes of multi-transmitter motor neurons in the abdominal ganglion that innervate the male Drosophila's internal reproductive organs (octopamine/glutamate and serotonin/glutamate), mapping the motor architecture that precisely times ejaculation during copulation, and showing the serotonergic class is required for male fertility while the octopaminergic class is not.