VISUAL ADAPTATION AND RETINAL GAIN CONTROLS
7
o
5 O
o
-%
~3 • o
E
2 A
z .~_
c9
1 •-~
z I I I I I
-5 -2 -1 0 1 2 2
8
LOG MEAN LUMINANCE ( c d / m Z )
FIG. 29. Contrast gain in cat retinal ganglion cells as a
function of mean luminance. The stimuli were drifting sine
gratings, presented at (Rayleigh) contrasts from 0.02 (2%)
up to 0.64 (64%). The temporal frequency of the drift was
4 Hz. Mean stimulus luminance was varied over a four to
five log unit range with neutral density f'dters. The CRT screen
had a white P4 phosphor. The photopic luminance of the
screen was measured; the scotopic equivalent luminance is
approximately twice the photopic for this light. Artificial
pupils, 3 mm in diameter, were used. With such a pupil, 1 cd
m -2 produces a retinal illumination of approximately 4.106
quanta(507 nm) (deg 2 s)-' on the cat's retina. The slope of
the linear portion of the response - contrast curve provided
an estimate of the contrast gain in impulses s-1 [contrast]-L
These are the results from three different on-centre X cells;
off-center and Y cells gave essentially similar results. The
responses at the highest mean luminance used, 100 cd m -s,
were cone-driven, but responses at lower mean levels were
rod-driven. From Shapley et al. (1983).
is t h a t o f E n r o t h - C u g e l l a n d R o b s o n ( 1 9 6 6 ) w h o
found a two-fold reduction in the receptive field
c e n t e r d i a m e t e r o f a n X cell w h e n t h e m e a n level
w a s v a r i e d f r o m 5" 10 .4 t o 16 c d m -2, a p p a r e n t l y
most of the change taking place at the low end of
t h e r a n g e o f m e a n levels, as s e e n i n Fig. 31. S i m i l a r
results have recently been obtained by Derrington
a n d L e n n i e ( 1 9 8 2 ) w h o r e p o r t a 32070 r e d u c t i o n i n
center diameter when the average illumination was
i n c r e a s e d f r o m 2 . 1 0 -3 t o 2 0 0 c d m - L S i n c e t h e s e
m e a n levels s p a n t h e r o d - c o n e t r a n s i t i o n i n t h e c a t
w h i c h o c c u r s a r o u n d 1 0 - 100 c d m -2 i n w h i t e l i g h t
( w h e n a 3 m m p u p i l d i a m e t e r is u s e d ; cf. E n r o t h -
C u g e l l et al., 1 9 7 7 a ) , o n e w o u l d w i s h t o h a v e m o r e
d e t a i l e d k n o w l e d g e a b o u t h o w t h e c e n t e r size v a r i e d
w i t h a v e r a g e l i g h t level b e f o r e f o r m u l a t i n g a n
explanation of the center's contraction in the light.
H o w e v e r , t h e r e s u l t s o f E n r o t h - C u g e l l et al. ( 1 9 7 7 b )
o n t h e r e l a t i v e sizes o f t h e r e c e p t i v e f i e l d c e n t e r
305
6 ~ i . e
|
p~
1,e~
2"6
1
0 t t I I I I I
0.1 0.2 0-3 O.S 0.7 1 2 3
Btimu]us c~meter, degrees
FIG. 30. Area-sensitivity curves at several background
luminances in cat ganglion cells. In this figure the label on
the ordinate, "sensitivity", is used to mean what we have
called "gain". The diameter of a circular spot centered on
the receptive field was varied in steps and the luminance
required to produce a criterion response was measured at each
value of the diameter. The temporal modulation was a 4 Hz
sinewave. The criterion response was just-audible
synchronization of the cell's discharge with the stimulus
modulation. The sloping and flat portions of these curves
intersect at a diameter denoted D, which is taken to be the
diameter of the central summing area of the receptive field
center. In these experiments the value of D, was
approximately constant across background level. Zero on the
"Relative sensitivity" scale corresponds to a stimulus
amplitude modulation of approximately 109 quanta(507 nm)
(deg 2 s)-' retinal illumination. The stimulus depth of
modulation was 0.6. The background was a circular spot with
a diameter of 8.5 deg. The retinal illuminations of the
background are given in log td, but these are "cat td", and
1 " c a t t d " is equivalent to approximately 6.105
quanta(507 nm) (deg2 s)-' on the retina. Thus the lowest
background illumination used in these experiments was
approximately 3.6. los quanta(deg 2 s)-1 on the retina. The
luminance required, through the 4 mm diam artificial pupil
used, to produce this lowest background illumination was
5.10-4 cd m 2. The brightest background was l0 s brighter. This
was an on-center cell, presumably a Y-cell. From Cleland
and Enroth-Cugell (1968).
w h e n it is d r i v e n b y r o d s o r d r i v e n b y c o n e s s u g g e s t
that the transition from rods to cones does not
p r o d u c e t h e c e n t e r ' s c o n t r a c t i o n . R a t h e r , it a p p e a r s
m o r e l i k e l y t h a t t h e c e n t e r size is r e l a t i v e l y c o n s t a n t
f r o m m i d - s c o t o p i c t o m i d - p h o t o p i c levels, a n d o n l y
becomes larger in the low scotopic range. This
i n f e r e n c e s h o u l d b e t e s t e d f u r t h e r s i n c e it m a y b e