Do Not: 5. Gain Control in Photoreceptors

324 R. M. SHAPLEYAND C.
ENROTH-CUGELL
personal communication). Figure 46 indicates that saturation (Dowling and Ripps, 1971). There is not
horizontal cells, and by inference the cones, of the one story for all vertebrates. The proximity of
catfish retina do not adapt but merely saturate, but horizontal cells to photoreceptors suggests that the
that bipolar cells, amacrine, and ganglion cells do receptors might also be very diverse in how they deal
adapt. In the catfish, much of the adaptation to with changes in mean or background illumination.
steady light must take place between cones and This expected diversity is found.
bipolar cells, though Fig. 46 also implies that there
are additional stages of gain control in the inner
plexiform layer. 5. GAIN C O N T R O L IN P H O T O R E C E P T O R S
There are data on the dependence of gain on
background in cat horizontal cells, under conditions Some photoreceptors adjust their gain by
such that the responses were due to photoreceptor adapting in the presence of steady illumination and
input from rods only. The graph in Fig. 49 indicates only saturate in very bright light, and other
that cat horizontal cells, driven by rods, have a gain photoreceptors adapt very little before they
vs background dependence which approximately saturate. This diversity of receptor function with
follows Weber's Law above 1 td [Note this is a "cat respect to gain control and saturation has been
t d " and therefore is equivalent to about 8.10 s suggested already in the discussion of gain controls
quanta(507 nm) (deg 2 s)-'; Steinberg, 1971]. Note in horizontal cells. In order to organize this diverse
that these horizontal cells typically receive mixed material, we will present the data on receptors as
photoreceptor input (Steinberg, 1971), even though follows: (a) photoreceptors which adapt a lot a n d
their major direct synaptic contact is with cones (see also saturate; and (b) photoreceptors which adapt
Appendix 1). This suggests that rod - cone coupling very little and mainly saturate.
is indeed important for determining the response
properties of horizontal cells. In comparing
horizontal cell responses with ganglion cell
4O
responses (Fig. 24), one notices that horizontal cell
gain does not begin to drop until the background
55
is 1 td, which is two to four log units higher than
the transition level of illumination for ganglion
.~_ 30
cells. This suggests that in the cat, as in the catfish, o_
there must be gain control mechanisms more
~ 25
proximal in the retina than the rods or the m
horizontal cells. However, the Weber Law behavior

~20
of horizontal ceils in the cat suggests that the rods
<1
may also adapt when the level of illumination is high l ii
o~ P5
enough (but see Section 5). Gain versus background
curves are not available for cone-driven horizontal
I0
cell responses in cat.
The diversity of horizontal cell gain changes due
05 - 1
to backgrounds indicated so far is representative of 05 0 0.5 ~0 r5 20 25 30
that seen generally. For instance, in m u d p u p p y the Log l~z, ,~ ( s c o l : o p i c t d )
horizontal cells behave as if influenced by saturation

and adaptation in the scotopic range, but resemble FIG. 49. Log test illumination (at 440 nm) plotted as a
turtle horizontal cells - - almost pure adaptation - - function of log background illumination (620 nm) for a
constant amplitude peak horizontal cell response
in the photopic range (Normann and Werblin, intracellularly recorded in the cat retina. The background
1974). However, carp horizontal cells show only duration was 5.5 - 6.5 s; the stimulus was 0.5 s in duration
saturation in the photopic range (Witkovsky, 1967). applied after the response to the background had settled to
a constant value. The stimulus was a spot on a larger
But the rod-driven skate horizontal cells do adapt background. The criterion response was 2.25 mV, and was
to light by gain reduction after an initial period of rod-driven. From Steinberg (1971).

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324 R. M. SHAPLEYAND C.

horizontal cells. However, the Weber Law behavior

that seen generally. For instance, in m u d p u p p y the Log l~z, ,~ ( s c o l : o p i c t d )

horizontal cells behave as if influenced by saturation

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