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Illusion - Is Seeing Really Believing?
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00048_Field_frep23.txt
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1996-12-30
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Around 1950, Stephen
Kuffler became the first to
record the responses of retinal
ganglion cells to spots of light
in a mammal, the cat. He was
then working at the Wilmer
Institute of Ophthalmology at
the Johns Hopkins Hospital. In
retrospect, his choice of
animals was lucky because the
cat's retina seems to have
neither the complexity of
movement responses we find in
the frog or rabbit retina nor the
color complications we find in
the retinas of fish, birds, or
monkeys. Kuffler used an
optical stimulator designed by
Samuel Talbot. This optical
device, a modified eye doctor's
ophthalmoscope, made it
possible to flood the retina with
steady, weak, uniform
background light and also to
project small, more intense
stimulus spots, while directly
observing both the stimulus and
the electrode tip. The
background light made it
possible to stimulate either rods
or cones or both, because only
the cones work when the
prevailing illumination is very
bright, and only the rods work
in very dim light. Kuffler
recorded extracellularly from
electrodes inserted through the
sclera (white of the eye)
directly into the retina from
the front. He had little
difficulty finding retinal
ganglion cells, which are just
under the surface and are fairly
large.
With a steady, diffuse
background light, or even in
utter darkness, most retinal
ganglion cells kept up a steady,
somewhat irregular firing of
impulses, at rates of from 1 to 2
up to about 20 impulses per
second. Because one might have
expected the cells to be silent
in complete darkness, this
firing itself came as a surprise.
By searching with a small
spot of light, Kuffler was able to
find a region in the retina
through which he could
influence--increase or
suppress--the retinal ganglion
cell's firing. This region was
the ganglion cell's receptive
field. As you might expect, the
receptive field was generally
centered at or very near the tip
of the electrode. It soon became
clear that ganglion cells were
of two types, and for reasons
that I will soon explain, he
called them on-center cells and
off-center cells. An on-center
cell discharged at a markedly
increased rate when a small
spot was turned on anywhere
within a well-defined area in
or near the center of the
receptive field. If you listen to
the discharges of such a cell
over a loudspeaker, you will
first hear spontaneous firing,
perhaps an occasional click,
and then, when the light goes
on, you will hear a barrage of
impulses that sounds like a
machine gun firing. We call
this form of response an on
response. When Kuffler moved
the spot of light a small
distance away from the center
of the receptive field, he
discovered that the light
suppressed the spontaneous
firing of the cell, and that
when he turned off the light
the cell gave a brisk burst of
impulses, lasting about 1
second. We call this entire
sequence--suppression during
light and discharge following
light--an off response.
Exploration of the receptive
field soon showed that it was
cleanly subdivided into a
circular on region surrounded
by a much larger ring-shaped
off region.