VISUALADAPTATIONANDRETINALGAIN CONTROLS
they produce " o n - off" responses to spots or full-
field flashes. The type N cells are like the so-called
"sustained" amacrine cells found by Kaneko (1973)
and Naka and Ohtsuka (1975). The change of gain
of these two types of cell with background is
illustrated in Fig. 46 from Naka et al. (1979). They
clearly are affected similarly by backgrounds and
also greatly resemble the ganglion cells in the way
they adapt. That is, the gain is approximately the
reciprocal of background, approximately Weber's
Law.
We have to make a brief digression to make clear
that the results in Figs 45 and 46 were obtained in
completely different ways. Werblin and
Copenhagen (1974) used standard rectangular
increments of light on a steady background, and
measured the peak of the change in intracellularly
recorded membrane potential. Naka et al. (1979)
measured first-order Wiener kernels by cross-
~./-- THEORETICALIO4
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"~"\ HORJZONTAL
z
~ 103 o,
~ 102 \XX~, IO2 I I I data. Clearly, the bipolars' gain begins to drop at
I00 I01 102
b
OFF CENTER
~BIPOLAR CELL
I--
_~ iO +
fTYPE N CELL
" G A N G LCI O~N
ioo
BIPOLARCELL
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MEAN INTENSITY
FIG. 46. Incremental gain o f catfish neurons, recorded
intraeellularly. The ordinate is the incremental gain, the
amplitude of the first order Wiener kernel normalized by the
magnitude o f the white-noise modulation. The horizontal
coordinate is the m e a n illumination. The dotted line is a
prediction from the Naka - R u s h t o n relation for the gain if
there were no adaptation, just saturation. Inset is an average
o f ten simultaneous recordings from horizontal and ganglion
cells. Results for various cell types are labeled in the figure.
We believe the lowest m e a n intensity was a retinal irradiance
of 0.1 ~ watts crn-2 on the retina. The light source was in most
c a s e s a g l o w - m o d u l a t o r tube which was attenuated by
neutral density filters. F r o m N a k a et al. (1979).
321
correlating a white-noise modulated light stimulus
with the resulting "noisy" modulation of the cells'
membrane potential. The gain for the type N cells,
and for all the other cells the results of which are
shown in Fig. 46, is the gain at the peak of the first
order Wiener kernel. The peak value of the kernel
(first or second order) at each background level,
divided by the standard deviation of the white noise
stimulus, may be taken as a measure of gain at that
background level.
4.2. Bipalar Cells
The bipolar cells form the main link between
photoreceptors and ganglion cells and thus the
dependence of gain on background for these cells
is of crucial importance for understanding the site
of adaptation.
The available data on gain control in bipolars
come mainly from Naka et al. (1979) and from
Thibos and Werblin (1978a). The results of Naka
et al. (1979) on catfish bipolars are displayed in
Fig. 46, on the same graph as the amacrine cell
about the same level of illumination as the
amacrines, and falls with a similar but somewhat
shallower slope. Since bipolar cells are the input to
amacrine cells in catfish (Naka and Ohtsuka, 1975)
as in other animals (Dowling and Boycott, 1965),
it is reasonable to suppose that the dependence of
gain on background observed in the amacrines is
already largely determined at the bipolar level.
The results on light adaptation in bipolars in
mudpuppy (Thibos and Werblin, 1978a) have
concentrated on the way in which the surround of
the bipolar receptive field sets the gain of the center.
This is illustrated in Fig. 47, which shows response
vs log illumination under two conditions: (i) no
background illumination; (ii) background
illumination falls on the bipolar's surround. In case
(i), the i n t e n s i t y - r e s p o n s e function is
approximately fit by the Naka - Rushton relation,
equation (6). In case (ii), the response vs
illumination curve of the same form is shifted to