320 R.M.
ENROTH-CUGELL
Y cells, due to substantial amacrine input to these
cells, are the basis for the psychophysical
" o n - o f f " transient sensitivity losses.
While the above results indicate the rapidity of
some gain control processes in the retina, they do
not rule out the existence of slower gain controls.
These slower processes have not been studied with
the Crawford paradigm, but they should be.
Adelson's (1982) results on the time course of
human light adaptation indicate the presence of
slower adaptation processes than have been studied
in ganglion cells up to now. It is well known but
poorly documented that when the mean level of
illumination is stepped up or down by two or more
log units in an adaptation experiment, the gain may
take several minutes to settle down or up to its new
steady value. Such slow adaptation processes
require further research.
4. GAIN AS A FUNCTION OF
I L L U M I N A T I O N IN A M A C R I N E CELLS,
B I P O L A R CELLS, AND H O R I Z O N T A L
CELLS
The phenomena of gain control in ganglion cells
require an explanation in terms of the functional
connections of the retinal network a n d / o r the
intrinsic p r o p e r t i e s o f the r e c e p t o r s and
interneurons in the retina. The fundamental
question is, where does the control of gain begin?
milllvolts
Io
8 5.0
/ /
4 - / I
I /
I /
2 - I I
I
0 ~ ' =l ' I
Lo 2.0 3.0
LOG I
Flo. 45. Curve-shifting in mudpuppy amacrine cells, recorded intracellularly. Diffuse illumination was used as a test
stimulus. The dashed curves are inferred in regions where no data were taken. The peak value of the response is plotted
against log flash illumination. The retinal illumination from the tungsten source used was about one quantum per rod per
second for log I = 0. This works out to roughly 6" 103 quanta (522 nm) (mm 2 s)-~ for log I = 0. From Werblin and Copenhagen
(1974).
SHAPLEY AND C.
Ideally, one would want to have the answer to this
question for the cat retina from which so much of
the results on gain control in ganglion cells have
been obtained. While there are some fragmentary
results on the cat, the technical difficulties of
i n t r a c e l l u l a r r e c o r d i n g have p r e v e n t e d a
comprehensive study of gain control in cat retinal
interneurons. We will therefore concentrate on the
results from the retinas of two cold-blooded
vertebrates which have been studied the most: the
mudpuppy (Normann and Werblin, 1974; Werblin,
1974; Werblin and Copenhagen, 1974; Thibos and
Werblin, 1978a, b) and the channel catfish (Naka
et al., 1979).
4.1. Amacrine Cells
In the mudpuppy the amacrine cells have a steep
response versus illumination curve which can be
shifted along the log illumination axis by steady
background illumination. This is illustrated in Fig.
45 from the work of Werblin and Copenhagen
(1974). This "curve shifting" is evidence for a gain
control at or prior to the amacrine cell (see Section
1.2.2.).
In the catfish, Naka et al. (1975) divided the
amacrine cells into two classes, the type N and type
C cells. The type C cells correspond to what Werblin
and colleagues (Werblin and Dowling, 1969;
Werblin and Copenhagen, 1974) call amacrine cells;
~.~ 4.5 ~-
.s
"-"~-~-~- 2.5 ~"
~ z . o
~"--,.~- ---,.5
J I t n
4.0 .... J [ ] Io mv
STIMULUS
Is