VISUAL ADAPTATION AND RETINAL GAIN CONTROLS 319

Between zero and 200 ms, the response declines
uniformly in magnitude. In related experiments, it
was shown that the gain stayed at the same new
value from 200 ms until 5 s after the conditioning
light was turned on. These experiments were done
in Y ceils under low scotopic conditions.
Similar results have been obtained in X cells of
the cat, by Saito and Fukada (1975), who studied
the time course of gain adjustment in X and Y cells
under mesopic or low photopic conditions. There
is a gap in our knowledge about the time course of
gain adjustment in X cells under scotopic
conditions, and a question about whether the time
course of adaptation changes markedly between
scotopic and photopic levels.
Saito and Fukada (1975) found that Y cells
showed a much sharper, more transient, gain
reduction immediately after the conditioning light
was turned on, and then again a reduction
immediately after it was extinguished, as indicated
in Fig. 44. This was at higher levels of background
XON
illumination than the earlier experiments of Enroth-
Cugell and Shapley (1973a), which may explain the
differences in findings. The gain reduction at both
" o n " and " o f f " of the conditioning light seen by
Saito and Fukada may possibly have been due to
one of the " a m a c r i n e " gain controls we have
postulated, either the contrast gaincontrol or the
steady-state flux gaincontrol of the Y cell center.
It seems more likely that the contrast gaincontrol
is involved in this phenomenon. The contrast
gaincontrol contains the kind of nonlinearity which
would cause reduction at both " o n " and " o f f " ;
it is an even-order nonlinearity which generates
responses of the same sign at " o n " and " o f f " ,
responses like those seen in amacrine cells and Y
cells. It is interesting that similar gain reductions
at " o n " and " o f f " are seen psychophysically in the
original Crawford (1947) experiment and in
intraretinal recording from the amacrine cell layer
(Gordon and Graham, 1973; also unpublished
results). It may be that transient gain reductions in
YON

Co) (e)

(c)

k .... I

(d) • |(h)
1.0
• S ' - -
I-0] ° f-
/. /

0.5- i

[ ! i i
I 2 o i 2 3
0 i 2 3 4 0 J 2 3

Time, sec Time, sec

FIG. 44. Crawford experiment for X and Y cells under low photopic or high scotopic conditions. X cell data are in (a) - (d);
Y cell data are ( e ) - (h). PST histograms show averaged responses to the test spot (short bar under histograms indicates
stimulus time) and to the conditioning spot (longer bar under histograms). The background luminance was 5.9 cd m -2 through
a 4 m m diameter artificial pupil. The unattenuated stimuli were : test, 3.9" 103, and conditioning, 6.8-10 ~ cd m "2. They
each subtended 5' diameter. The test spot was placed in the middle of the receptive field center, while the conditioning
spot was placed 15' to the side, still within the center. (a) and (e): the test spot was presented 2 s after onset o f the conditioning
spot. (b) and (f): test spot alone, as a control. (c) and (g): test spot presented 2 s after offset o f the conditioning spot.
The duration of the test spot was 200 ms for the X cell, 100 ms for the Y cell. The duration o f the conditioning spot w a s
3 s. The vertical calibration is 50 impulses s-'. (d) and (h): The time courses o f the change of magnitude o f the response
to the test caused by presentation of the conditioning spot. The relative magnitude of the response is plotted against the
delay between the onset times o f test and conditioning spots. From Saito and Fukada (1975).

PRR3-1*