318 R. M . SHAPLEY AND C.
ENROTH-CUGELL
test stimuli by Enroth-Cugell and Jakiela (1980).
The effect of the contrast gaincontrol on square
wave responses is to make them smaller and more
transient, as can be seen in Fig. 42 from Enroth-
Cugell and Jakiela (1980). This is an example of
nonlinear spatial summation because the peripheral
stimulus generates no response when presented
alone but suppresses the response to the centrally
placed bar stimulus when both are presented
together. This kind of suppressive effect from
peripheral stimulation has been observed in many
ganglion cell types in several different species.
ON-CENTERY-CELL
-.~uI001
50] grating
(a)
~ 0
~ 100
drifting
~ 50]
L
(b)
o
t (c)
~ ( d )
0 l 2 3 sec
FIG. 42. Reduction of the center's gain by a moving pattern
in the periphery of the receptive field in a cat retinal ganglion
cell. Responses of an on-center Y cell to a centered bar, 1 deg
wide, 13 deg high, of luminance 0.25 cd m "2 flashing at
0.4 Hz.
(a) The bar was superimposed on a background of
luminance 5 c d m -~. Both bar and background were generated
on a C R T which subtended 13 deg by 16 deg.
(b) A drifting sine grating with a contrast of 0.5, and spatial
frequency 1.25 c deg -1 covered the C R T screen except for
a 2 deg wide band down the center of the screen where the
stimulus bar was presented. The grating was drifting with
a temporal frequency of 1.14 Hz.
(c) Shows the time course of the bar stimulus.
(d) Indicates the time course of the luminance change at
a point in the field over which the grating was drifting. From
Enroth-Cugell and Jakiela (1980).
3.9. T i m e C o u r s e o f G a i n A d j u s t m e n t in R e t i n a l
Ganglion Cells
The retinal gain as measured in ganglion cells is
readjusted rapidly but not instantaneously by
sudden changes in background level. This has been
determined in physiological experiments which are
analogous to Crawford's psychophysical experi-
ment on the time course of light adaptation in
humans (Crawford, 1947).
=c ¶
o22:
210
N 0.6
~02
~ I 1 I I I 1 I [ t
}- 0 1O0 300 500 700
Delay between 'on" of conditioning
and ' o n ' of test light (rnsec)
FtG. 43. Time course of gain change in cat ganglion cells:
modified Crawford experiment. On a 12 deg steady
background of retinal illumination 5' los quanta(507 nm)
(deg 2 s)-1 were centered: (a) a square wave modulated
conditioning spot (1.5 s on, 3.5 s off), diam. 0.57 deg, retinal
illumination 2.5. los quanta(deg 2 s)-'; (b) a test spot, flashed
on for 20 ms in one cell (filled circles) and for 50 ms in
another cell (empty circles), of diam. 0.1 deg, with retinal
illumination 3. l0 s quanta(507 nm) (deg ~ s)-'. The test stimuli
were presented with varying delays with respect to the onset
of the conditioning light. Plotted in the figure for two
different on-center cells are the magnitudes of the responses
to the test pulses of light at different delay times between
onset of conditioning light and onset of test pulse. The arrows
point to the responses to the test stimuli which generated
responses which coincided in time with the peak of the
response to the conditioning light. From Enroth-Cugell and
Shapley (1973a).
In this experiment retinal gain is probed by
measurement of the magnitude of the response to
a brief test pulse of light. The test pulse is applied
in the dark and at various times after a conditioning
light is turned on. The results of an experiment of
this sort are shown in Fig. 43 from Enroth-Cugell
and Shapley (1973a). The response to the test pulse
of light is the same a half second after the
conditioning light is off as it is when the
conditioning light has been off for several minutes.
The response is definitely reduced within 200 ms
from the time the conditioning light is turned on.