324 R. M. SHAPLEYAND C.
ENROTH-CUGELL
personal communication). Figure 46 indicates that
horizontal cells, and by inference the cones, of the
catfish retina do not adapt but merely saturate, but
that bipolar cells, amacrine, and ganglion cells do
adapt. In the catfish, much of the adaptation to
steady light must take place between cones and
bipolar cells, though Fig. 46 also implies that there
are additional stages of gain control in the inner
plexiform layer.
There are data on the dependence of gain on
background in cat horizontal cells, under conditions
such that the responses were due to photoreceptor
input from rods only. The graph in Fig. 49 indicates
that cat horizontal cells, driven by rods, have a gain
vs background dependence which approximately
follows Weber's Law above 1 td [Note this is a "cat
t d " and therefore is equivalent to about 8.10 s
quanta(507 nm) (deg 2 s)-'; Steinberg, 1971]. Note
that these horizontal cells typically receive mixed
photoreceptor input (Steinberg, 1971), even though
their major direct synaptic contact is with cones (see
Appendix 1). This suggests that rod - cone coupling
is indeed important for determining the response
properties of horizontal cells. In comparing
horizontal cell responses with ganglion cell
responses (Fig. 24), one notices that horizontal cell
gain does not begin to drop until the background
is 1 td, which is two to four log units higher than
the transition level of illumination for ganglion
cells. This suggests that in the cat, as in the catfish,
there must be gain control mechanisms more
proximal in the retina than the rods or the
horizontal cells. However, the Weber Law behavior
of horizontal ceils in the cat suggests that the rods
may also adapt when the level of illumination is high
enough (but see Section 5). Gain versus background
curves are not available for cone-driven horizontal
cell responses in cat.
The diversity of horizontal cell gain changes due
to backgrounds indicated so far is representative of
that seen generally. For instance, in m u d p u p p y the
horizontal cells behave as if influenced by saturation
and adaptation in the scotopic range, but resemble
turtle horizontal cells - - almost pure adaptation - -
in the photopic range (Normann and Werblin,
1974). However, carp horizontal cells show only
saturation in the photopic range (Witkovsky, 1967).
But the rod-driven skate horizontal cells do adapt
to light by gain reduction after an initial period of
saturation (Dowling and Ripps, 1971). There is not
one story for all vertebrates. The proximity of
horizontal cells to photoreceptors suggests that the
receptors might also be very diverse in how they deal
with changes in mean or background illumination.
This expected diversity is found.
5. GAIN C O N T R O L IN P H O T O R E C E P T O R S
Some photoreceptors adjust their gain by
adapting in the presence of steady illumination and
only saturate in very bright light, and other
photoreceptors adapt very little before they
saturate. This diversity of receptor function with
respect to gain control and saturation has been
suggested already in the discussion of gain controls
in horizontal cells. In order to organize this diverse
material, we will present the data on receptors as
follows: (a) photoreceptors which adapt a lot a n d
also saturate; and (b) photoreceptors which adapt
very little and mainly saturate.
4O
55
.~_ 30
o_
~ 25
m
~20
<1
l ii
o~ P5
I0
05 - 1
05 0 0.5 ~0 r5 20 25 30
Log l~z, ,~ ( s c o l : o p i c t d )
FIG. 49. Log test illumination (at 440 nm) plotted as a
function of log background illumination (620 nm) for a
constant amplitude peak horizontal cell response
intracellularly recorded in the cat retina. The background
duration was 5.5 - 6.5 s; the stimulus was 0.5 s in duration
applied after the response to the background had settled to
a constant value. The stimulus was a spot on a larger
background. The criterion response was 2.25 mV, and was
rod-driven. From Steinberg (1971).