268 R.M. SHAPLEY AND C.
ENROTH-CUGELL
or n-fold brighter, so then is the retinal image of the white
paper 2-,3-, or n-fold brighter, but so also is the rest of
the visual field and the entire retina receives the 2-,3-, or
n-fold illumination. The ratio of the quantity of light on
the entire retina and the image of the paper remains
constant under otherwise equal conditions, l think,
therefore, that a process is initiated whose intensity
depends on this ratio, and which causes the sensation of
white for the retinal image. The brightness of the retinal
image is, so to speak, being evaluated in terms of the total
excitation. This is a judgement, the psychological side of
the matter. The physical side is the process mentioned.
It has not yet been discovered."
Mach's example shows a deep insight into the
perception of brightness and the purpose of visual
adaptation. However, there are some details of his
brief analysis which are not quite right. In
appreciating what is still valid and what needs
correction, one may begin to see the point of light
adaptation.
We will discuss two related but distinct comments
in the quoted paragraph. The first is, "White of a
lesser intensity appears gray next to a brighter
white". This observation can be explained by
modern psychophysics which has established that
the brightness of an object is determined, to a great
extent, by the Weber contrast between the object
and its surroundings (Heinemann, 1955, 1972;
Whittle and Challands, 1969). However, it is also
known that brightness is mainly determined by the
contrast near the border between an object and its
surroundings. This surprising conclusion is forced
by two experiments. First, the brightness difference
between the two regions of unequal luminance
varies directly with the sharpness of the border
between them, being maximal for the steepest
border (Thomas and Kovar, 1965; Shapley and
Tolhurst, 1973). Second, two regions of equal
luminance appear of unequal brightness when a
local luminance difference is introduced as a border
between them (Ratliff, 1965; Craik, 1966;
Cornsweet, 1970; Land and McCann, 1971; Shapley
and Tolhurst, 1973). See Fig. 2 for an illustration
of this effect. Thus, the dependence of brightness
on contrast, which Mach referred to in his 1865
paper, is now known to be mainly a dependence on
border contrast.
The relation between brightness, contrast, and
light adaptation emerges from a critical examination
of a second remark made by Mach, which concerns
brightness constancy, " . . . we are never in doubt
whether we have before us a white or gray paper
even u n d e r quite d i f f e r e n t c o n d i t i o n s o f
illumination...". This observation can be
explained by the fact that the contrast of an object
on a background is not changed by variation in the
level of illumination. Thus, if the visual system has
the ability to derive brightness from contrast, it will
thereby achieve brightness constancy. We will at this
point demonstrate that contrast is invariant with
illumination, and then show how the calculation of
contrast by the visual system can be explained in
terms of retinal adaptation.
The invariance of contrast with changes in the
level of illumination can be demonstrated by an
example. Consider as the simplest case a uniformly
illuminated scene with an object on a background.
The situation is illustrated in Fig. 3. The light
coming from the object side of the border is
proportional to I. Ro where I is the light falling on
the scene from the source of illumination, and Ro is
the reflectance of the object. The light reflected from
the background side of the border is I . R B. The
Weber contrast is (IRo-IRB)/IR B. Dividing numer-
ator and denominator by I yields (Ro-RB)/RB as the
contrast of an object of reflectance Ro upon a
background of reflectance Ra. Thus, the contrast
is independent of the level o f illumination I and
depends only on the reflectances of object and back-
ground.
Now we must show that retinal adaptation
provides the mechanism by which the visual system
responds to contrast. Consider what happens when
the receptive field of a retinal cell (see Appendix 2)
crosses the border between an object and a
background, as in the example of Fig. 3. Suppose
that the receptive field is " l o o k i n g " at the
background just before an eye movement occurs
[Fig. 3(a)], and that the eye movement causes the
receptive field to cross the border [Fig. 3(b)]. The
change in the amount of light falling on the
receptive field of the neural unit is the stimulus
which elicits a neural signal which identifies the
border. The stimulus is thus IRo-IR w Now we
must consider the role of adaptation. A neuron
crossing the border only has been " l o o k i n g " at the
background side of the border. So the neuron is
adapted to IR~. As a reasonable hypothesis about
what adaptation does, to be justified by data later,
we propose it adjusts the gain of the neuron to be