290 R. M. SHAPLEY AND C.
ENROTH-CUGELL
If the gain of the cell were related to this steady state
response, the gain reduction would be less for the
large disk. However, physiological results on
s c o t o p i c a d a p t a t i o n do not s u p p o r t this
interpretation of the psychophysical result (Enroth-
Cugell et al., 1975; Barlow and Levick, 1976). That
is, the gain of the receptive field center in cat retinal
ganglion cells in the scotopic range decreases and
then levels off as background diameter is increased,
and is not a decreasing and then increasing function
as would be supposed from the conventional
interpretation of sensitization. F u r t h e r m o r e ,
psychophysical sensitization in the scotopic range
has not been convincingly demonstrated under
stabilized image conditions (MacLeod, 1978). This
suggests that retinal or central stimulation by the
moving border of the background disk may be
needed for sensitization to be observed (MacLeod,
1978). Moreover, the relation of sensitization to
retinal adaptation has been questioned by Lennie
and MacLeod (1973) who showed that the key to
sensitization was the uniformity in luminance of the
0.75 deg diameter desensitizing disk and the outer
annulus. If the outer annulus were either lower or
higher in luminance than the central disk it would
reduce sensitization.
2.2. The Cone System
2.2.1. SENSITIVITY AS A FUNCTION OF BACKGROUND
The cone system (Photopic system) is like the rod
system in yielding Weber's Law under some
circumstances and the square root law under others.
As is evident from Fig. 8 from Wyszecki and Stiles
(1967), in human vision the cones take over at
threshold from the rods above 0.1 to 1 td in
background retinal illumination. The increment
threshold curves in the literature tend to all show
a cone plateau (cone " d a r k light"; Barlow, 1958)
from 0.1 up to about 10 td. The photopic Weber's
Law for a moderate-sized test spot (diameter > 0.5
deg) takes over from the cone " d a r k light" limited
behavior at 10 td very reliably.
Although the photopic thresholds for spots
follow Weber's Law up to the point where the
pigment is bleached away, it has recently been
shown that saturation can be demonstrated in the
cone system with adapting backgrounds which are
themselves small in area (Buss et al., 1982). This
saturation can begin at as low a background as
10 td, in the low photopic range. On a small
diameter (0.25 deg) background the threshold of a
small spot climbs much more steeply than Weber's
Law (obtained with a large background) would
predict. This effect may be observed under
stabilized image conditions (Tulunay-Keesey and
Vassilev, 1974). The photopic saturation is thought
to be due to saturation of a cone-driven interneuron
in the retina, one which has a center-surround
interaction so that a background of large area will
release the putative interneuron from saturation.
Electrophysiological evidence on sensitization in the
photopic range is presently scant and equivocal, but
there are some indications it may exist (see Section
3.7.).
2.2.2. CONTRAST SENSITIVITY AND MEAN LEVEL
For most targets, and in particular moderate-
sized spots with a sharp edge on a large background,
Weber's Law holds from 10 td to l0 s td, i.e.
throughout the photopic range of backgrounds.
This is shown very clearly in the data of Whittle and
Challands (1969), Fig. 17. The lower curve for each
subject is the increment threshold curve; above it
are curves of constant brightness as matched to a
test flash in the contralateral eye. It is interesting
that the c o n s t a n t b r i g h t n e s s curves are
approximately parallel to the increment threshold
curve. This means that the apparent brightness of
two different increments was proportional to their
(Weber) contrasts. Thus, Whittle and Challands'
results imply that (Weber) contrast rather than
luminance determined the apparent brightness.
They also point out that other, central, factors may
contribute to apparent brightness. Under the
conditions of their experiments, the mechanisms
which determined brightness were purely monocular
and therefore probably retinal - - as evidenced by
the fact that stimulus contrast against a fairly high
background in one eye was needed to match a much
weaker flash luminance against zero background in
the other eye. These data of Whittle and Challands
support the main hypotheses we started out with
about the functional significance of adaptation in
establishing brightness constancy contingent on
contrast, which contrast depends on the reflectances