READMElc4.md

LC4

systemVisionspeciesD. melanogaster3D view25 of 626 cellssources5

LC4 is a lobula columnar visual projection neuron type of the fruit fly that responds to looming, the rapid expansion of an approaching object’s image. LC4 neurons synapse directly onto the giant fiber escape neuron and convey how fast the object expands.

Also known asLobula columnar type 4 · ColA (in larger flies)
RegionLobula; axons end in the LC4 optic glomerulus, where they meet the giant fiber
SystemVision: eye and optic lobe
In the display bundle25 of the 626 reconstructed neurons in the site’s MaleCNS display bundle, which shows 626 of the model’s 166,700 neurons: LC4 25

Where they sit

LC4 dendrites lie in the lobula and their axons converge in the LC4 optic glomerulus, which is innervated by the giant fiber, designated DNp01 (Namiki et al., 2018).

What they do

Removing this input to the giant fiber during looming stimuli showed that it encodes the angular expansion velocity of the approaching object, while other inputs encode its angular size; the giant fiber integrates the two features linearly (von Reyn et al., 2017). The size component comes from LPLC2, and electron microscopy shows that both LC4 and LPLC2 synapse directly onto the giant fiber (Ache et al., 2019).

LC4 neurons viewing different parts of the visual field make opposite, graded numbers of synapses onto two descending neurons, DNp02 and DNp11, along the front-to-back axis of visual space. Activating DNp11 makes flies lean forward and DNp02 promotes backward takeoffs, so the location of a threat is converted into the direction of escape (Dombrovski et al., 2023).

Behavior

Optogenetic activation of LC4 drives jumping (Wu et al., 2016). Fast looms bias the fly toward rapid, giant-fiber-mediated escapes (von Reyn et al., 2017).

3D view: LC4

MaleCNS v1.0 · 25 of 626 reconstructed neurons

The 3D view needs JavaScript and WebGL.

25 of 626 reconstructed neurons in this display bundle

Highlighted: the LC4 cells among the 626 reconstructed neurons of the site’s display bundle, which shows 626 of the 166,700 neurons in the MaleCNS v1.0 model with skeletons simplified for the web. Anatomy only: a highlight marks cell identity, not activity. Anatomy: MaleCNS v1.0, FlyEM / HHMI Janelia, CC BY 4.0.

In TheFlyBrain

In TheFlyBrain experiments

Several experiments measure LC4 together with LPLC2 as a looming readout, and some can drive LC4 and LPLC2 directly through an engineered encoder; direct drive bypasses retinal detection.

8 experiments name these cells in their setup or measurement notes. On the workstation, what the fly sees reaches the fixed 166,700-neuron model and spikes are recorded from cells like these.

  • 0001Media ObservationTimed cues stimulate LC10a and LC4/LPLC2 through the engineered movie-cues-v1 encoder. Over 700 ms of model time, mean DNa02 rate plus 0.25 x DNp01 rate is compared with a threshold to request a board visit.
  • 0003Retinal BenchChannels: retinal input, looming feature cells (LC4, LPLC2) and moving-target cells (LC10a); side both, left or right; amplitude 0–1.2.
  • 0045Torture Chamber: Six Documented DevicesDefault input is retina only. Optional direct contact/load drives the station's annotated tactile, chordotonal or campaniform populations; direct loom/escape drives LC4/LPLC2 and DNp01, at amplitude 0–1.2.
  • 0053The Ludovico TechniqueThe panel reports measured spikes for LC4 + LPLC2 (looming), LC10a (small target), HS + H2 (horizontal motion), DNa02 (steering) and DNp01 (giant fiber).
  • 0056The Star GateMeasured readouts during playback: horizontal motion (HS + H2), vertical motion (VS) and looming (LC4 + LPLC2).
  • 0070Looming Escape (Fly Swatter)Each 200 ms window carries ten 20 ms retinal frames, so the loom expands step by step; measured readouts are DNp01 and LC4 + LPLC2.
  • 0071Ganzfeld ProtocolDuring live playback the pick is the total measured spikes in four readout groups (LC10a, LC4 + LPLC2, HS + H2, DNa02) during judging, modulo 4; otherwise it is the seeded, pre-programmed pick.
  • 0072DreamachineMeasured readouts: LC4 + LPLC2, DNp01, HS + H2 and VS. Input is retinal only; weights are fixed.

Sources

  1. Namiki S, Dickinson MH, Wong AM, Korff W, Card GM (2018). The functional organization of descending sensory-motor pathways in Drosophila. eLife 7:e34272. doi:10.7554/eLife.34272
  2. von Reyn CR, Nern A, Williamson WR, et al. (2017). Feature Integration Drives Probabilistic Behavior in the Drosophila Escape Response. Neuron 94:1190–1204.e6. doi:10.1016/j.neuron.2017.05.036
  3. Ache JM, Polsky J, Alghailani S, et al. (2019). Neural Basis for Looming Size and Velocity Encoding in the Drosophila Giant Fiber Escape Pathway. Current Biology 29:1073–1081.e4. doi:10.1016/j.cub.2019.01.079
  4. Dombrovski M, Peek MY, Park JY, et al. (2023). Synaptic gradients transform object location to action. Nature 613:534–542. doi:10.1038/s41586-022-05562-8
  5. Wu M, Nern A, Williamson WR, et al. (2016). Visual projection neurons in the Drosophila lobula link feature detection to distinct behavioral programs. eLife 5:e21022. doi:10.7554/eLife.21022

Anatomy in the 3D view: MaleCNS v1.0, FlyEM / HHMI Janelia, CC BY 4.0. Summaries on this page describe published findings in living flies; the workstation’s model is a fixed-weight spiking model of the wiring diagram.

Glossary: Looming · Visual projection neuron · Lobula · Descending neuron