Retinal Sampling
Neural Sampling of the Retinal Image
Retinal Architecture
• Neural processing of the retinal image begins with the
transduction of light energy into corresponding changes of
membrane potential of individual light-sensitive photoreceptor
cells called rods and cones.
• Photoreceptors are laid out across the retina as a thin sheet that
varies systematically in composition as shown schematically
following figure.
• At the very center of the foveal region, which corresponds roughly
to the center of the eye's field of view, the photoreceptors are
exclusively cones.
• Because cone-based vision is not as sensitive as rod-based vision,
the central fovea is blind to dim lights (e.g., faint stars) that are not
clearly visible when viewed indirectly.
• At a radial distance of about 0.1-0.2 mm along the retinal surface
(0.35-0.7° field angle) from the foveal center, rods first appear and
in the peripheral retina rods are far more numerous than cones.
• Although rod and cone diameters grow slightly with distance from
the fovea, the most dramatic change in neural organization is the
increase in spacing between cones and the filling in of gaps by large
numbers of rods.
• Consequently, cones occupy only about 30% of the retinal surface
and the density of rods is about 30 times that of cones.
• Given this arrangement of the photoreceptor mosaic, we may
characterize the first neural stage of the visual system as a sampling
process by which a continuous optical image on the retina is
transduced by two interdigitated arrays of sampling apertures.
• The cone array supports photopic (daylight) vision and the rod
array supports scotopic (night) vision. In either case, the result is a
discrete array of neural signals called a neural image.
• The neural images encoded by the rod and cone mosaics are
transmitted from eye to brain over an optic nerve which, in
humans, contains roughly one million individual fibers per eye.
• Each fiber is an outgrowth of a third-order retinal neuron called a
ganglion cell.
• It is a general feature of the vertebrate retina that ganglion cells
are functionally connected to many rods and cones by means of
intermediate, second-order neurons called bipolar cells.
• As a result, a given ganglion cell typically responds to light
falling over a relatively large receptive field covering numerous
rods and cones.
• Neighboring ganglion cells may receive input from the same
receptor, which implies that ganglion cell receptive fields may
physically overlap.
• Thus in general the mapping from photoreceptors to optic nerve
fibers is both many-to-one and one-to-many.
• The net result, however, is a significant degree of image
compression since the human eye contains about five times more
cones, and about 100 times more rods than optic nerve fibers.
• In humans and other primates, one particular class of retinal
ganglion cell called P-cells (by physiologists) or midget cells (by
anatomists) is by far the most numerous everywhere across the
retina (Rodieck, 1988).
• This P-system has evolved to meet the perceptual requirements for
high spatial resolution by minimizing the degree of convergence
from cones onto ganglion cells.
• Ultimate limit to this evolutionary strategy is achieved in fovea
where individual cones exclusively drive not just one, but two
ganglion cells via separate interneurons (bipolar cells).
• In an eye of an individual with normal color vision, the cone
population that drives the P-system consists of two subtypes (L-
and M- type) with slightly different spectral sensitivities.
• Since a foveal ganglion cell is functionally connected to a single
cone, the ganglion cell will inherit the cone's spectral selectivity,
thereby preserving chromatic signals necessary for color vision.
• In peripheral retina, P-ganglion cells may pool signals
indiscriminately from different cone types, thus diminishing our
ability to distinguish colors.
Functional Implications of Neural Sampling
1. Contrast detection and the size of sampling elements
The neural architecture of the retina outlined above
has important functional implications for the basic
visual tasks of contrast detection and spatial
resolution.
• The finest spatial pattern for which contrast is detectable depends
ultimately upon the size of the largest receptive fields in the chain
of neurons that supports contrast perception.
• Since ganglion cell receptive fields can be no smaller than that of
individual cones, and are generally expected to be larger, the retinal
limit imposed on contrast detection will be set by the spatial
filtering characteristics of ganglion cell receptive fields.
2. Spatial resolution and the spacing of sampling elements
For two points to be discriminated, at least one relatively
unstimulated photoreceptor must lie between two relatively
stimulated photoreceptors.
Although this rule was formulated in the context of
resolving two points of light, it applies equally well to the
case of resolving the individual bars of a sinusoidal grating
as illustrated in figure below
• Although the neural image is meant to be a faithful
representation of the retinal image, these two kinds of
images are fundamentally different: the optical image
formed on the retina is spatially continuous, whereas the
neural image is discrete.
• This difference between the stimulus and its neural
representation is central to the sampling theory of visual
resolution, and it raises an important theoretical question.
• Is information lost by converting a continuous light image
into a discrete neural image?
• In fact, given the proper method of interpolation,
Shannon's rigorous sampling theorem states that
“the envelope will exactly reconstruct the retinal image
provided that the sampling process is error-free and that
the sampling density is sufficient to provide at least two
samples per cycle of the highest spatial frequency
component in the image.”
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