VISUAL ADAPTATION AND RETINAL GAIN CONTROLS
ON-CENTER OFF-CENTER
RECEPTIVE FIELD
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FIG. 64. Schematic representation of the mutual antagonism
between center and surround in on and off-center cells. In
on-center cells (a,b,c), an increment in the center increases
the cell's firing rate while an increment of light in the surround
decreases the firing rate. When the two increments are applied
together in synchrony, as in (c), the response to light " o n "
is less than in (a) and the response to light " o f f " is less than
in (b). This means the two regions are mutually antagonistic.
In off-center cells (d,e,f), the cells' response pattern at light
" o n " in the center and in the periphery are reversed, but the
mutual antagonism is the same.
responses to increments of light, the surround
would cause inhibitory responses to increments. In
off-center cells in which the central region was
inhibitory during an increment, the surround would
be excitatory during an increment. The on- and off-
center cells and their center-surround organization
are illustrated in Fig. 64.
Rodieck (1965) made a major advance by
developing a model for the cat ganglion cell
receptive field in terms of overlapping center and
surround mechanisms. Each mechanism may be
conceived of as the receptors and interneurons, the
signals of which are pooled together to influence
the firing of the ganglion cell. Within the center
mechanism all light evoked signals generated within
the pool of these receptors and interneurons are
summed, according to the model, and similarly for
the surround. Then center and surround signals are
summed at the ganglion cell. The signals from
different positions within each pool are weighted
by what Rodieck called the "sensitivity profile",
and which we will refer to as the "spatial
distribution of gain". The center has its own narrow
spatial distribution of gain, and the surround has
a rather broader spatial distribution. Rodieck
341
proposed that these two spatial distributions could
be approximated by Gaussian surfaces with
different extents of spread. The spatial resolution
and optimal spatial tuning of retinal ganglion cells
can be rationalized in terms of Rodieck's model
(Enroth-Cugell and Robson, 1966). The spatial
resolution of the cell is due to the finite size of the
I center, and in fact can be predicted from knowledge
I of the magnitude of the spread of the center's
Gaussian spatial distribution of gain (Cleland et eL,
1979; So and Shapley, 1979; Linsenmeier et el.,
1982). Figure 65 shows the Rodieck model for the
receptive field of cat retinal ganglion cells.
~ R-MECHANISM
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CENTER TYPE
RESPONSE
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FIG. 65. The two spatially overlapping mechanisms in the
Rodieck model of retinal ganglion cell receptive fields. In
the sketch at the top, the horizontal axis represents distance
on the retina. The heights of the two curves represent the
gain of the center and o f the surround, as labeled, as a
function o f position on retina. Both are Gaussian functions
of position; the center's Gaussian has a narrower spread than
the surround's. The center and surround have opposite sign
in this model. This results in mutual antagonism. In this
model the center and surround components combine by
simple addition, i.e. linearly. Thus, response to stimulation
anywhere within the field is, according to the model, simply
a s u m of the center and surround components in response
to the stimulus, as is illustrated in the figure. From Rodieck
(1973).