VISUAL ADAPTATION AND RETINAL GAIN CONTROLS 307

3ifi
0.1 02 03 0 5 0.7 I 2
Diameter of adapting spot,degrees
3
' :fb rj00I 02 03 0507 I
StimuLus d i a m e t e r , degrees
2 3

F[6. 32. Comparison of adaptive (a) and signal (b) summation areas of an on-center cat ganglion cell.
(a) The horizontal axis gives the log of the diameter of unmodulated adapting spots. The vertical axis gives the log of
the reciprocal of the adapting spot illumination in relative units. The adapting illumination was set so that the cell produced
a criterion response to a small (0.13 deg) centrally located spot of fixed luminance, which was sinusoidally modulated (4 Hz,
0.6 contrast).
(b) The horizontal axis givesthe log of the diameterof the stimulus, which was again a spot modulated at 4 Hz, sinusoidally,
at 0.6 contrast. But in this experiment the stimulus diameter was varied. The vertical axis in (10)gives the log of the relative
gain for the spots of different size; it is also the log of the reciprocal of the illumination required to elicit a constant response.
In 0a), the spots were presented on a steady background of 6.10 ~quanta(dega s)-'. The criterion response in both (a) and
(b) was just audible synchronization of the cell's discharge with the stimulus modulation. From Cleland and Enroth-Cugell
(1968).

stopped. The important result of this experiment
is that the signal summation area and the adaptive
summation area were the same. Probably most if
not all of these experiments were on Y retinal
ganglion cells, but later work indicates similar
results hold for X cells (Harding, 1977). The results
are the same for on- and off-cells. Very similar
results have been obtained with this experimental
design on rat optic tract fibers by Green et al. (1977)
and Tong and Green (1977). Results of this kind
have also been obtained in lower vertebrates: in frog
ganglion cells (Reuter, 1969; Burkhardt and
Berntson, 1972), and in goldfish ganglion cells
(Schellart and Spekreijse, 1972).
Related experiments by Shapley et al. (1972) and
by Enroth-Cugell and Shapley (1973b) indicate that,
as the area of an adapting spot of fixed luminance
is increased, the gain declines. This is illustrated in
Fig. 33. The (fixed) luminance of the adapting spot
was chosen so that for the smallest adapting spot
the gain had not been reduced from its dark adapted
maximum. As the area of the adapting spot was
increased, gain declined and the response to the test
stimulus became more transient. Results similar to
these were obtained by Schellart and Spekreijse
(1972) in the goldfish retina. They found that the
temporal impulse response was speeded up and
made more diphasic by increasing the area of a
background spot of constant luminance.
Another experiment which demonstrates the
significance of the area as well as the luminance of
the background is illustrated in Fig. 34. The figure
shows the actual responses of a ganglion cell to
turning on adapting spots of quite different area
and luminances. The luminance of the larger one
had been adjusted until it produced the same gain
reduction as the smaller adapting spot, as indicated
by the response to the superimposed brief test flash.
Equal adaptive effect is associated with equal
responses of the ganglion cell to the " a d a p t i n g "
spots. The conclusion from all these experiments
is that the gain of the ganglion cell center
mechanism is set by the sum of all the steady state
input to the center. This in turn implies that the
adaptive effect of a background on the center
mechanism of a ganglion cell is, under the
conditions of these experiments, determined by the
total effective flux: the sum of all the light per unit
time which falls on the center from the background,
weighted by the spatial "sensitivity profile"
(Cleland and Enroth-Cugell, 1968; Enroth-Cugell
and Shapley, 1973b).
The meaning of these results in a psychophysical
context is that the summation pool of a ganglion
cell is the same size as the adaptation pool. The
meaning in a neuroanatomical context is that
whatever interneuron determines the size of the
receptive field center is also implicated in the gain
control process.
In the light o f these results on the dependence of