The sector of the ONH that is related to the location of a visual stimulus as it hits the retina is governed by the path that axons from the stimulated retinal ganglion cells (RGCs) take to exit the eye. Although much is known about the growth of RGC axons in utero, the exact mechanism is not well understood.
10 It is known that the axons grow from the RGCs across the retina and exit the eye through the ONH and that they tend to fasciculate together to form small bundles.
10 If we make the assumption that when axons develop they take the shortest path from the RGC to the ONH and that the eye is a sphere, we can immediately generate a location-to-sector map. Unfortunately, such a map does not bear much relation to our observations of the RNFL in the retina, as nearly all locations in the nasal visual field (both inferior and superior) are mapped to sector 1 of the G map in
Figure 1 .
Not only would this lead to a large number of axons tracking across the fovea, we know from RNFL thickness measurements of normal eyes
11 that the RNFL is thin at the temporal margin of the disc, relative to the superior and inferior areas. Hence, having more than half the RGCs underlying visual field locations with their axons exiting the eye in sector 1 of the G map is clearly incorrect. If this were the case, then the temporal margin would be the thickest part of the RNFL profile around the optic disc. We can alter the model to take this into account by imposing an order in which RGCs grow their axons, and then only allowing an axon into a certain sector if that sector is not already “full” of other axons according to the RNFL thickness profile around the disc. This also has the side effect of fascicularization: The axons tend to form bundles.
Figure 2describes a simple algorithm (dubbed GROW) for realizing this model. To summarize, the ONH is divided into sectors, and the retina is divided into a grid of elements. The RGCs in each element attempt to assign their axons to the sector of the ONH that is closest along the surface of a sphere. If the sector is already “full” of axons, according to an RNFL thickness profile, then an alternate sector must be found. The order in which elements are chosen to grow their axons therefore is important, as elements chosen early in the process are likely to find room in their closest sector, whereas later elements are likely to have to find new sectors. Although the algorithm does not explicitly prohibit axons from tracking across the foveal pit, if elements are ordered according to their distance from a point midway between the fovea and the ONH, this effect happens automatically.
Information of the distribution of RGCs on the retina required for steps 1 and 5 in
Figure 2are taken from the data published by Curcio and Allen.
12 The profile of RNFL thickness required in step 2 is taken from
3 Figure 4in Hood and Kardon,
13 simply normalized to sum to 1 and assuming that on average the thickness of axons is roughly the same in each sector. In step 3, for the experiments reported herein, we use a hexagonal grid with each hexagon having a side length of 0.2 mm, and shorten the distance of each hexagon from the start point by a factor of 1.28 in the horizontal axis. This is based on the observation of Curcio and Allen
12 that the contours of RGC density follow an elliptical pattern, with the horizontal axis of the ellipse being 1.28 longer than the vertical on average.
12 We chose the starting point of the growth to be slightly to the ONH side of the fovea (−2.5°). The starting point is based in part on the knowledge that central RGCs grow their axons before peripheral RGCs,
10 and in part by the observation that RGCs close to the foveal side of the ONH would surely have axons that track directly into the ONH, and not in some arcuate path. This second claim is supported by the G map, where RGCs in sector 1 are all between the fovea and the ONH. We fix the position of the ONH at (−15°, 0°), and discuss possible ramifications of this in the final section of the paper. Finally, if steps 12 and 13 of GROW are reached—that is, the sector on the shortest path is full, and an alternate sector must be found—we search sectors in an order that respects the horizontal meridian. For example, if we are constructing a map with the six sectors of the G map and sector 6 is full, we try sector 5; and if sector 2 is full, we try sector 3. Although such a scheme does not explicitly prevent RGCs from having axons that cross the horizontal midline to enter the ONH, with the parameters chosen in this article, such a crossing does not occur. Also note that we are assuming that each RGC has a single axon entering the ONH, and so
\[{{\sum}_{e}}\ C_{e}{=}\ {{\sum}_{i}}\ S_{i}{=}R\]
where the parameters for this equation are defined in
Figure 2 .