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318 R. M . SHAPLEY AND C.

ENROTH-CUGELL

test stimuli by 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


The effect of the contrast gaincontrol on square Ganglion Cells
wave responses is to make them smaller and more
transient, as can be seen in Fig. 42 from Enroth- The retinal gain as measured in ganglion cells is
Cugell and Jakiela (1980). This is an example of readjusted rapidly but not instantaneously by
nonlinear spatial summation because the peripheral sudden changes in background level. This has been
stimulus generates no response when presented determined in physiological experiments which are
alone but suppresses the response to the centrally analogous to Crawford's psychophysical experi-
placed bar stimulus when both are presented ment on the time course of light adaptation in
together. This kind of suppressive effect from humans (Crawford, 1947).
peripheral stimulation has been observed in many
ganglion cell types in several different species. =c ¶
o22:

ON-CENTERY-CELL
210

N 0.6

-.~uI001 ~02
~ I 1 I I I 1 I [ t
50] grating }- 0 1O0 300 500 700
Delay between 'on" of conditioning
(a) and ' o n ' of test light (rnsec)
~ 0
FtG. 43. Time course of gain change in cat ganglion cells:
~ 100 modified Crawford experiment. On a 12 deg steady
drifting background of retinal illumination 5' los quanta(507 nm)
(deg 2 s)-1 were centered: (a) a square wave modulated
~ 50] conditioning spot (1.5 s on, 3.5 s off), diam. 0.57 deg, retinal
L
illumination 2.5. los quanta(deg 2 s)-'; (b) a test spot, flashed
(b) on for 20 ms in one cell (filled circles) and for 50 ms in
o
another cell (empty circles), of diam. 0.1 deg, with retinal
illumination 3. l0 s quanta(507 nm) (deg ~ s)-'. The test stimuli
t (c) were presented with varying delays with respect to the onset
of the conditioning light. Plotted in the figure for two
~ ( d ) 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
0 l 2 3 sec 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).
FIG. 42. Reduction of the center's gain by a moving pattern
in the periphery of the receptive field in a cat retinal ganglion In this experiment retinal gain is probed by
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 measurement of the magnitude of the response to
0.4 Hz. a brief test pulse of light. The test pulse is applied
(a) The bar was superimposed on a background of in the dark and at various times after a conditioning
luminance 5 c d m -~. Both bar and background were generated
on a C R T which subtended 13 deg by 16 deg. light is turned on. The results of an experiment of
(b) A drifting sine grating with a contrast of 0.5, and spatial this sort are shown in Fig. 43 from Enroth-Cugell
frequency 1.25 c deg -1 covered the C R T screen except for and Shapley (1973a). The response to the test pulse
a 2 deg wide band down the center of the screen where the
stimulus bar was presented. The grating was drifting with of light is the same a half second after the
a temporal frequency of 1.14 Hz. conditioning light is off as it is when the
(c) Shows the time course of the bar stimulus. conditioning light has been off for several minutes.
(d) Indicates the time course of the luminance change at
a point in the field over which the grating was drifting. From The response is definitely reduced within 200 ms
Enroth-Cugell and Jakiela (1980). from the time the conditioning light is turned on.