VISUAL ADAPTATION AND RETINAL GAIN CONTROLS
Recently, Linsenmeier et al. (1982) have taken a
fresh look at this question by measuring the gain
of X and Y ganglion cells in the cat in response to
drifting gratings on high scotopic or mesopic
backgrounds. They also gauged the size of the
center of each ganglion cell by fitting the observed
dependence of contrast gain on spatial frequency
with a "Difference of Gaussians" model. The
spatial sensitivity profiles of receptive field center
and surround are approximated by Gaussian
functions in this model. The spatial spread of the
center's Gaussian is a measure of the effective
radius of the center's distribution of sensitivity (or,
more precisely, gain). Figure 37 is their graph of
the peak gain of the center plotted vs the center's
effective radius, for a large population of cat retinal
ganglion cells. The figure demonstrates that the cells
with the largest centers had the lowest peak gain,
and that the gain was approximately the inverse of
the center's radius. While this result is qualitatively
like the earlier results of Enroth-Cugell and
Shapley, Fischer and May, and Cleland et al., it is
quantitatively different in that gain in the light
adapted state is inverse to the radius and not the
area of the center. However, there is quite a lot of
variance of gain across the population of ganglion
cells, so much so that Linsenmeier et al.'s results
do not conclusively disprove the a r e a - g a i n
relation. Furthermore, since their measurements
were made at backgrounds which might be in the
high scotopic or in the mesopic range, the precise
value of the slope o f the gain vs area line might be
influenced by the degree to which rods or cones are
the predominant photoreceptor input for cells of
different sizes. These qualifying remarks suggest
that the book is not closed on the dependence of
gain on area of the receptive field center. As
suggested below, the hypothesis of Enroth-Cugell
and Shapley (1973b), that gain varies inversely with
center area in the light-adapted state, is useful in
rationalizing p s y c h o p h y s i c a l results on the
dependence of sensitivity vs background curves on
target size.
The interpretation of these area effects in
adaptation must be modified by the discovery that
there is not a wide variation in receptive field center
size among ganglion cells of one type, X or Y, at
a given retinal locus (Cleland et al., 1979; So and
Shapley, 1979). The coefficient of variation of the
313
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FiG. 37. Relation between the center's size and its peak gain
in the middle of the receptive field. The dependence of
(Rayleigh)contrast gain on spatial frequencywas determined
for X and Y cells by adjustment of contrast to reach a
constant response criterion. The experimental curves were
fit with a Difference of Gaussians model as in Fig. 31, from
which both the center's radius and its gain at the peak of
its sensitivity profile could be determined. These two values
are plotted against each other to show that, at the same mean
luminance, cells with larger centers have lower gain. The
empty symbolsare for on-centercells, the filled symbolsare
for off-center cells. Circles denote X cells; triangles denote
Y cells. The pupillaryarea was 16 mmL The mean luminance
was around 14 cd m-L From Linsenmeier et al. (1982).
center-diameter distribution at any one retinal locus
is at most 0.25 (So and Shapley, 1979) and is
probably less in an individual animal. There is a
marked increase in receptive field center-diameter
at retinal loci away from the area centralis; the
d i a m e t e r o f the center is a p p r o x i m a t e l y
proportional to the distance from area centralis.
This is true for both X and Y cells, and for both
on- and off-center cells. On- and off-cells have
approximately the same size at any one locus on the
retina. As stated above, Y cells have an
approximately ten times larger area than X cells at
each locus. The combination of these facts with the
preceding results on the effects of area on
adaptation leads to the following conclusions. First,
cells with larger receptive fields in the periphery of
the retina ought to be more light-adapted than
central ganglion cells with smaller centers, under
conditions of uniform constant b a c k g r o u n d
illumination. Second, Y ganglion cells ought to be
more light-adapted than X ganglion cells at the same
retinal locus. By the degree of light-adaptation we
mean the degree to which gain has been reduced