READMEr1-r6-photoreceptors.md

R1–R6 photoreceptors

systemVisionspeciesD. melanogaster3D view16 of 626 cellssources6

R1–R6 are the six outer photoreceptor cells in each ommatidium, or unit eye, of the fruit fly’s compound eye. They all belong to one spectral class and are the main input to the fly’s motion vision.

Also known asOuter photoreceptors · R1-6
RegionCompound eye (retina); axons end in the lamina
SystemVision: eye and optic lobe
In the display bundle16 of the 626 reconstructed neurons in the site’s MaleCNS display bundle, which shows 626 of the model’s 166,700 neurons: R1-6 16

What they are

Each ommatidium contains eight photoreceptors, R1 to R8 (Meinertzhagen and O'Neil, 1991). R1–R6 are arranged around the outside of the ommatidium and R7 and R8 in its centre. Whereas R7 and R8 come in several spectral classes, R1–R6 all belong to a single one (Wardill et al., 2012).

Light is absorbed by rhodopsin in the rhabdomere, a stack of membrane folds, and converted into an electrical signal by a cascade that is the fastest G-protein signalling pathway known (Hardie and Raghu, 2001). Like other fly photoreceptors, R1–R6 respond with graded changes in membrane voltage rather than spikes.

Where they sit

Their cell bodies lie in the retina and their axons end in the lamina, the first neuropil of the optic lobe. There R1–R6 terminals provide the photoreceptor input to the lamina’s repeating modules, the cartridges, and synapse onto the lamina monopolar cells (Meinertzhagen and O'Neil, 1991). R1–R6 cells in six neighbouring ommatidia that look at the same point in space send their axons to the same cartridge, an arrangement called neural superposition.

What they do

Genetic dissection of the lamina places R1–R6 at the start of the achromatic motion channel: their signal passes mainly through lamina neurons L1 and L2, which together are necessary and largely sufficient for motion-dependent behavior (Rister et al., 2007). Downstream, the pathway splits into ON and OFF channels that end in the T4 and T5 motion detectors (Joesch et al., 2010; Maisak et al., 2013).

R1–R6 are not the only route: R7 and R8 also feed the motion pathway, which broadens its spectral tuning and improves motion discrimination (Wardill et al., 2012).

Behavior

Turning responses to moving patterns depend on motion signals that begin in R1–R6 and pass through L1 and L2 (Rister et al., 2007; Maisak et al., 2013).

3D view: R1–R6

MaleCNS v1.0 · 16 of 626 reconstructed neurons

The 3D view needs JavaScript and WebGL.

16 of 626 reconstructed neurons in this display bundle

Highlighted: the R1–R6 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 the workstation, each scene is rendered from the fly’s head and averaged into 32 luminance columns, which drive the model’s R1–R6, R7 and R8 cells by estimated azimuth. The input has no color or vertical structure and is not a model of compound-eye optics. See retinal input.

One experiment names 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 ObservationEvery 250 ms, television pixels or a render from the fly's head are reduced to 32 luminance columns that drive the model's photoreceptors.

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. Hardie RC, Raghu P (2001). Visual transduction in Drosophila. Nature 413:186–193. doi:10.1038/35093002
  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. 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
  5. Maisak MS, Haag J, Ammer G, et al. (2013). A directional tuning map of Drosophila elementary motion detectors. Nature 500:212–216. doi:10.1038/nature12320
  6. Wardill TJ, List O, Li X, et al. (2012). Multiple Spectral Inputs Improve Motion Discrimination in the Drosophila Visual System. Science 336:925–931. doi:10.1126/science.1215317

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: Ommatidium · Compound eye · Photoreceptor · Retinal input (32 columns) · Lamina