312 R. M. SHAPLEY AND C.

ENROTH-CUGELL

the retina after center and surround have been ] l I I I I I

segregated. The neuroanatomy of the cat retina tells
us that all post-receptoral interneurons pool the
activity of m a n y photoreceptors. Therefore, the
p o s t - r e c e p t o r a l gain c o n t r o l m u s t p o o l
photoreceptor signals.
3.6. Gain Control and Receptive Field Size Across
the Population of Ganglion Cells
Since l i g h t - e v o k e d neural signals are summed
over the receptive field center to set the gain of the
center of a particular ganglion cell, one naturally
would guess that ganglion cells with centers of
different sizes would be light adapted to different
extents by a large uniform background. The results
on the spatial summation of adaptive effect suggest
that gain depends on the total, steady state, effective
flux, i.e. illumination multiplied by area weighted
by the center's gain per unit area. The total effective
flux falling on the center of a cell with a small
receptive field will be less than the flux falling on
a large receptive field center; the gain should be
reduced less in the small receptive field center.
The initial test of this idea is offered in Fig. 36,
from Enroth-Cugell and Shapley (1973b). What is
plotted in Fig. 36 is the transition illumination at
the knee of the curve relating gain and background
illumination. The transition illumination is defined
here empirically as the illumination at which the
gain has dropped by a factor of two from the dark
adapted gain (Note the slight difference between this
definition and the more rigorous definition of
transition illumination in connection with equation
(20)). Because the data in Fig. 36 were collected
from many different cats, possibly in different
physiological states, the transition illumination was
multiplied by the dark adapted gain to obtain a
corrected transition illumination. This corrected
transition illumination is plotted against center
summing area, determined from an area - threshold
curve (Cleland and Enroth-Cugell, 1968). It can be
seen in Fig. 36 that the cells with larger centers have
a lower effective transition level, and that the
transition level is approximately proportional to the
center summing area. These data might be
compatible with other functions of center size
besides area, because of the large variance. The cells
U
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10 ~ o ~ o o
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025 05 10 25 50 10 25
Total summing areaof centr resp mech (deg~)
FIG. 36. Transition level as a function of center area for a
population of cat retinal ganglion cells. The value of the
transition level from the horizontal to the sloping portions
of the gain vs background illumination curves (as in Fig. 24)
is plotted against center summingarea. Empty circlesare from
on-center cells, while filled circles are from off-center cells.
The cells were not grouped into X and Y classes. From
Enroth-Cugell and Shapley (1973b).
in this study were not classified as X or Y.
Therefore, a question unresolved by these results
is whether X and Y cells have the same dependence
of effective transition level on center area.
Further evidence on the dependence of gain
setting on receptive field center size across the pop-
ulation of ganglion cells comes from the concordant
studies of Fischer and May (1970) and Cleland et
al. (1973). The results of both studies implied that
the center's gain, defined as G = d R / d F where F
is luminous flux (illumination times area), is
inversely proportional to the center's summing area
when the ganglion cell is well light adapted in the
mid-scotopic to mesopic range by large uniform
backgrounds. This is consistent with the result of
Enroth-Cugell and Shapley (1973b) on the "effec-
tive transition level" and with the approximately
inverse relationship between gain and background
above the transition level, equation (21). Thus, three
studies seem consistent in supporting the hypothesis
that ganglion cells with larger centers are more light-
adapted than those with smaller centers under the
same fixed uniform background conditions, because
of spatial summation of adapting signals.