VISUAL ADAPTATION AND RETINAL GAIN CONTROLS
processes which change the magnitude of the noise
from which the signal must be picked out as well
as by those which affect the size of the signal.
In p h y s i o l o g i c a l e x p e r i m e n t s , the w o r d
"sensitivity" is usually used to mean the reciprocal
of the stimulus required to produce a neural
response of a criterion size. In this meaning of the
word, noise is ignored because neural responses are
usually averaged over a number of identical
stimulus cycles to eliminate noise as much as
possible. There has been confusion in the literature
when a result on psychophysical sensitivity has been
t a k e n to imply s o m e t h i n g d e f i n i t e a b o u t
physiological sensitivity or vice versa, because in one
case noise has an effect and in the other it does not.
We will try to avoid confusion by calling
psychophysical sensitivity, "'sensitivity", and
physiological sensitivity, "'gain". Gain we define as
the ratio of the magnitude of the physiological
response to the stimulus magnitude, in the small-
signal range in which response is proportional to
the stimulus. Gain thus has units like mV/quantum
in photoreceptors, or impulse/quantum in retinal
g a n g l i o n cells. A l t h o u g h we distinguish
physiological gain from psychophysical sensitivity,
the two are related. If the internal noise were more
or less unaffected by background light, a reduction
in retinal gain as a consequence of an increase in
background would produce a corresponding
reduction in visual sensitivity.
1.1.2. ADAPTATION
Considered in terms of sensitivity and gain, the
one unambiguous traditional term which describes
the adaptation state is total dark adaptation, the
state of highest sensitivity reached by an observer
and highest retinal gain reached by a living retina
that has been left in total darkness for a few hours.
We define light adaptation as those variations in
the properties of the visual system from the totally
dark adapted state which are produced by variation
in the level of light. For instance, light may change
the gain of the retina, or its time course of response,
or its spectral sensitivity, or its spatial summation
properties. However, an increase of retinal noise
due to the random times of arrivals of light quanta
is not strictly speaking a property of the retina
which is changed by light. But it is a factor which
could lead to an increase of the psychophysical
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threshold. Therefore, in order to understand how
light adaptation contributes to vision, specifically
to the variation of visual sensitivity with mean level
of illumination, one must distinguish between
adaptation and the effects of increased noise caused
by light. Similarly, as we will show, gain can be
reduced by saturation, e.g. the limitation on the
amplitude of response imposed by a response
ceiling. We wish to distinguish this kind of gain
reduction due to saturation, or as it has been called
"response compression", and the gain control of
adaptation which involves a change in the properties
of the retina with time during illumination.
1.1.3. BRIGHTNESS, LUMINANCE, AND RETINAL
ILLUMINATION
Throughout our discussion, brightness means
apparent brightness, the subjective sensation of how
light or dark an object is. The objective measure
of the amount o f light emanating from a luminous
source or reflecting object, weighted by the
observer's spectral sensitivity function, is called
luminance (Wyszecki and Stiles, 1967, p. 372). The
illumination falling onto a surface from a distant
luminous source is proportional to the luminance
of the source multiplied by the square of the
numerical aperture of the optical system between
source and surface. The numerical aperture of an
optical system is the reciprocal of its f-number. For
the eye, the numerical aperture is proportional to
the pupil diameter, and the retinal illumination is
therefore proportional to the area of the pupil.
Luminance can be expressed in terms of effective
quanta of light per unit solid angle per unit time
per unit area of the source. Illumination can be
expressed in terms of effective quanta of light per
unit time per unit area of the surface on which the
light is falling. We will write L for stimulus
luminance and I for retinal illumination, with the
understanding that in most experiments the pupil
area is fixed and therefore the two quantities are
simply proportional. Evidence will be presented
below about the importance o f the luminous flux
of light falling on a receptive field. Flux is
illumination multiplied by area and can be expressed
in units of effective quanta per unit time.