302 R. M. SHAPLEY AND C.

ENROTH-CUGELL

3.2. Gain Control in the Photopic Range
The inputs from cones to cat ganglion cells can
be isolated from those from rods by means of Stiles'
two-color technique (see Wyszecki and Stiles, 1967,
p. 572). When this is done, it is found that cone
signals have a constant gain over the range that the
gain of rod signals drops by a factor of one
thousand or more (Enroth-Cugell et al., 1977a),
from total dark adaptation to the high scotopic
range. This can be seen in Fig. 27. Plotted there are
the threshold illuminations for criterion cone-driven
and rod-driven responses to be elicited by a test
stimulus on a blue background, as a function of the
level of background. For a single ganglion cell one
obtains a two-branched gain vs background curve
which is reminiscent of the two-branched,
psychophysical sensitivity vs background curves
(Fig. 8). This indicates that the separate gain control
of rod and cone signals is achieved by the retina
prior to the ganglion cells, and that rod and cone
signals are kept segregated at least up to the points
at which the gains are set in the parallel " r o d " and
" c o n e " pathways through the retina. These results
are found for receptive field centers of both on- and
off-center cells, and for both X and Y cells. This
has the added implication that rod and cone signals
are segregated in both the X and Y pathways until
the gain is set.
The results in Fig. 27 imply independence of
adaptation mechanisms for the rod and cone
pathways to the ganglion cells in the cat. However,
Nelson (1977) found that rod signals are coupled
into cones. His work led him to the conclusion that
the main pathway of rod signals to horizontal cells
was through the cones. Presumably a similar
conclusion would apply to the bipolar cells. That
is, rod signals should travel through cone bipolar
cells because of the large amount of coupling of rod
signals into cones. This poses a problem, namely
how can the cone and rod signals adapt separately
when they are carried by the same interneurons?
One possible explanation is that all adaptation may
take place in receptors, but that explanation has

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FIG. 27. Gain for the rod and cone systems in a cat retinal ganglion cell. The illumination required to evoke a criterion
response is graphed vs the background illumination. The points plotted as filled symbols were obtained with a blue - green
stimulus light on a b l u e - green background, and were taken to represent the gain of the rod pathway. Their values a r e
given by their heights on the left vertical axis, in equivalent quanta of a monochromatic 507 nm light. The empty symbols
are the criterion illuminations for a red spot of light on the same blue - green background, and are interpreted as indicating
the cone pathway's gain. The response criterion was just-audible synchrony of the cell's firing rate with the 4 Hz square
wave modulation of the small stimulus spot (0.2 deg diameter), which was located in the middle of the receptive field.
In this figure, retinal illuminations are referred to quanta at the cornea, before losses in the eye. From Enroth-Cugell et
al. (1977a).