VISUAL ADAPTATION AND RETINAL GAIN CONTROLS 323

to a neural gain control acting on the bipolar and
not on the horizontal cell. But at higher levels of
mean illumination, the horizontal cell potential
starts saturating and its gain plummets, failing
below the bipolar's gain. Since bipolar cells appear
to have steeper intensity- response functions than
horizontal cells (Werblin, 1974), they might saturate
at even lower mean levels than the horizontal cells,
were it not for the saving action of the gain control.
The automatic gain control causes the bipolar cells
(and the more proximal retinal neurons they feed)
to lose some gain at lower levels in order to preserve
gain at higher levels which would otherwise be lost
because of saturation (see Section 1.2.2.).
4.3. Horizontal Cells
There is quite a lot of information on the control
of gain in horizontal cells because they are easier
to record from intracellularly than the other retinal
interneurons. Since horizontal cells are anatomically
one synapse away from photoreceptors, they often
mimic the receptors' adaptational properties. One
of the significant outcomes of this fact is that the
regulation of gain in photoreceptors may be
inferred, with caution, from studies of the
regulation by light of the gain of horizontal cells.
~/j" , I
• Dark adapted
• - 6 3 BG
-50 O 5.3 BG
• - 4 . 2 BG
-4O -32 BG
We can begin to see some of the diversity in
photoreceptor adaptation in the variety of
horizontal cells' adaptational behavior (cf. also
Section 5).
Perhaps the best studied receptor - horizontal cell
system is in the turtle retina. Results on adaptation
in turtle horizontal cells are quite clear, as shown
in Fig. 48 (Normann and Perlman, 1979b). The
intensity-response curves shift to the right with
background, and the gain follows Weber's Law.
These horizontal cells are driven exclusively by long-
wavelength cones, and their adaptational properties
are mainly determined by their cone inputs
(Normann and Perlman, 1979a). The curve-
shifting in Fig. 48 is clear evidence for adaptation
of the type suggested in equation (10), with an
automatic gain control located in the cones.
To illustrate inter-species diversity we compare
the turtle horizontal cell with the horizontal cell of
the catfish retina, the gain vs background curve of
which is illustrated in Fig. 46. Here the horizontal
cell curve is not like the curve of Weber's Law.
Rather, it follows the dashed curve, which is a
prediction based on the N a k a - R u s h t o n relation
[equations (6) and (7)]. That is, the loss of gain is
caused by saturation. Under the conditions of these
experiments, the catfish horizontal cells were driven
exclusively from long-wavelength cones (Naka,
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-7.0 -6.0 -5 0 -40 ~30 -20
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F{G. 48. Adaptation in turtle horizontal cells; intracellular recording. The stimuli were 0.5 s increments or decrements on
a background, except for the results on the dark adapted eye, which were increments only. The stimuli were large 3.2 m m
diameter spots on the retina. Peak responses measured from the dark-adapted resting potential (dotted line) were plotted
as a function of test illumination. The curves were drawn by eye. The horizontal bar through each curve indicates the steady
m e m b r a n e potential measured at least two minutes after background onset. The illuminations are given as log attenuation.
The unattenuated test stimulus (0 log) was 6.4-10 ~ quanta(640 nm) (cm 2 s)-1 on the retina. The unattenuated background
illumination was 9.1.10 TM quanta(640 nm) (cm 2 s)-1. From N o r m a n n and Perlman (1979b).