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284 R. M . SHAPLEY AND C.

ENROTH-CUGELL

I I I 1 I I
retinal ganglion cells with different-sized receptive
10
fields to make sense out of the transition from
square root law to Weber's law (Enroth-Cugell and
Shapley, 1973b). In essence, the explanation is that
the channels which respond to low spatial
frequencies suffer more loss of sensitivity with
-/
l I I I
ecc
L I
increase in background than do the channels which
respond to high spatial frequencies because retinal
0o 10 ° 50 ° gain controls depend on areal summation of
background illumination. The channels which are
FIG. 12. Lowest threshold modulation depth as a function most sensitive to low spatial frequencies, according
of retinal eccentricity and mean level of illumination. These
data are all from contrast threshold measurements for sine to this view, sum adaptive signals over a larger area
grating patterns which subtended 4 deg by 4 deg and were than do the channels most sensitive to high spatial
drifting at a temporal frequency of 4 Hz. The stimuli were frequencies. A more detailed explanation of this
presented on a Hewlett - Packard 1310A display with a P 15
phosphor (dominant wavelength at 510 nm), and were viewed point is offered in the Theory section.
by the subject through a 2 m m diameter artificial pupil.
Contrast sensitivity curves were measured at each eccentricity 2.1.4. HUMAN AND FELINE CONTRAST SENSITIVITIES
and illumination, and the m i n i m u m threshold modulation
depth (the reciprocal of the peak contrast sensitivity across COMPARED
spatial frequency) for each condition was denoted m*, which
Since most of the electrophysiological results on
is the quantity plotted in the figure. The different symbols
are for different mean illuminations. E m p t y circles are for retinal adaptation and gain control come from
10 td; filled circles are for 1 td; circles with dots in the center experiments on animals, it is relevant to compare
are for 0.1 td. From Koenderink et al. (1978).
h u m a n p s y c h o p h y s i c a l and cat b e h a v i o r a l
measurements of contrast sensitivity at different
mean levels of illumination. Recently, this has
sensitivity over the scotopic. It seems that the foveal
become possible because of the results of Pasternak
photopic contrast sensitivity is especially high when
and Merigan (1981). Their results show that there
compared with the photopic (and scotopic) contrast are noticeable differences between the contrast
sensitivity of the peripheral retina. They also found sensitivities of cat and man. As can be seen in
that, for a stimulus of a given size and spatial
Fig. 13, the cat's contrast sensitivity improves with
frequency, the transition from square root law to mean level in a way analogous to man, but at all
Weber's law depended on the retinal eccentricity; mean levels the cat's peak contrast sensitivity is
the transition occurred at lower luminances the lower. This may be because the high mean
more peripheral was the stimulus.
luminances were photopic for the human, but all
mean luminances were probably scotopic for the
2.1.3. THE CHANNELHYPOTHESIS
cat. The r o d - c o n e transition in cats is at a much
There exists a working hypothesis one can use to higher mean luminance than in man, presumably
interpret Barlow's, Blackwell's, Daitch and because of the much higher rod - cone ratio in the
Green's, and Koenderink et al.'s, findings about the cat (Lennie et al., 1976). Contrast sensitivity grows
influence of size or spatial frequency or retinal much more steeply for high spatial frequencies than
eccentricity on the transition from square root law for low, in cat as in man. However, the acuity of
to Weber's Law. This hypothesis is that the visual the cat is much worse than that of man, as can be
system is composed of a set of size or spatial seen in Fig. 13 by the separation between the high
frequency channels (Campbell and Robson, 1968; spatial frequency cutoffs of the contrast sensitivity
Enroth-Cugell and Robson, 1966; Robson, 1975; curves at all mean levels except the lowest.
Graham, 1980). It is supposed that a test stimulus However, the basic similarities between the
sifts through the channels and excites only those dependences of feline and h u m a n contrast
channels which are " t u n e d " to the stimulus. Such sensitivities on mean level of illumination encourage
an hypothesis can be used in conjunction with us to attempt a synthesis of human psychophysical
research findings on the adaptation properties of studies of light adaptation with the body of

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