READMElamina-monopolar-cells.md

Lamina monopolar cells L1–L5

systemVisionspeciesD. melanogaster3D view82 of 626 cellssources5

Lamina monopolar cells L1 to L5 are the output neurons of the lamina, the first neuropil of the fruit fly’s optic lobe. They relay photoreceptor signals to the medulla, where L1 and L2 begin the ON and OFF pathways for motion vision.

Also known asL1 · L2 · L3 · L4 · L5 · Lamina neurons
RegionLamina (optic lobe); axons end in the medulla
SystemVision: eye and optic lobe
In the display bundle82 of the 626 reconstructed neurons in the site’s MaleCNS display bundle, which shows 626 of the model’s 166,700 neurons: L1 24, L2 25, L3 22, L4 5, L5 6

What they are

The lamina is built from repeating modules called cartridges, one for each point in the visual field. A complete electron-microscopy reconstruction of a cartridge identified 12 neuron types: the R1–R6 photoreceptor terminals, the R7 and R8 fibres, the five monopolar cells L1–L5, the medulla neurons C2, C3 and T1, and a wide-field amacrine cell (Meinertzhagen and O'Neil, 1991).

Where they sit

L1–L5 have their branches in the lamina and send their axons to the medulla. At the lamina, the R1–R6 signal splits into four parallel pathways: L1, L2, L3 and the amacrine cell–T1 pathway (Rister et al., 2007).

What they do

Blocking L1 eliminates the responses of motion-sensitive neurons to moving bright edges, and blocking L2 eliminates responses to moving dark edges. Photoreceptor signals therefore segregate into an ON channel beginning with L1 and an OFF channel beginning with L2, much like ON and OFF bipolar cells in the vertebrate retina (Joesch et al., 2010).

L3 was first linked to orientation behavior rather than motion (Rister et al., 2007) and was later shown to act together with L1 and L2 as an input to motion-detecting circuits (Silies et al., 2013). A screen of all 12 lamina neuron classes found that L2 and L4, together with the feedback neurons C2 and C3, are required for normal detection of directional motion, and that nearly all lamina classes contribute (Tuthill et al., 2013).

Behavior

At high pattern contrast, the L1 and L2 pathways are redundant for motion-guided behavior; at intermediate contrast, L2 mediates front-to-back and L1 back-to-front motion (Rister et al., 2007).

3D view: L1–L5

MaleCNS v1.0 · 82 of 626 reconstructed neurons

The 3D view needs JavaScript and WebGL.

82 of 626 reconstructed neurons in this display bundle

Highlighted: the L1–L5 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

In a living fly, photoreceptors and lamina neurons signal with graded potentials rather than spikes. The workstation’s model represents them, like every other cell, as leaky integrate-and-fire units.

No experiment in the registry names these cells in its setup or measurement notes yet. They are part of the fixed 166,700-neuron MaleCNS model that every experiment runs on.

Sources

  1. Meinertzhagen IA, O'Neil SD (1991). Synaptic organization of columnar elements in the lamina of the wild type in Drosophila melanogaster. Journal of Comparative Neurology 305:232–263. doi:10.1002/cne.903050206
  2. Joesch M, Schnell B, Raghu SV, Reiff DF, Borst A (2010). ON and OFF pathways in Drosophila motion vision. Nature 468:300–304. doi:10.1038/nature09545
  3. Rister J, Pauls D, Schnell B, et al. (2007). Dissection of the Peripheral Motion Channel in the Visual System of Drosophila melanogaster. Neuron 56:155–170. doi:10.1016/j.neuron.2007.09.014
  4. Silies M, Gohl DM, Fisher YE, Freifeld L, Clark DA, Clandinin TR (2013). Modular Use of Peripheral Input Channels Tunes Motion-Detecting Circuitry. Neuron 79:111–127. doi:10.1016/j.neuron.2013.04.029
  5. Tuthill JC, Nern A, Holtz SL, Rubin GM, Reiser MB (2013). Contributions of the 12 Neuron Classes in the Fly Lamina to Motion Vision. Neuron 79:128–140. doi:10.1016/j.neuron.2013.05.024

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: Lamina · ON and OFF pathways · Optic lobe · Leaky integrate-and-fire (LIF)