264 R.M. SHAPLEY AND C.
ENROTH-CUGELL
1. INTRODUCTION
Vision is our primary sensory channel for
interaction with the outside world. It allows us to
recognize familiar faces and creatures, and objects;
it allows us to orient ourselves in space and to
navigate from place to place. It is a pathway for
esthetic e n j o y m e n t and for i n f o r m a t i o n
transmission. The visual system is one of the many
miracles of nature.
Intensive study of the visual process has revealed
that the retina must perform several operations on
the image delivered by the eye's optics in order to
make manageable the difficult jobs of the
brain : pattern recognition and spatial localization.
One of the basic operations the retina performs is
the subject of this chapter: retinal adaptation. The
retinal neurons adapt to variations in illumination
by changing their gain and response time course.
The purpose of adaptation is to keep the retinal
response to visual objects approximately the same
when the level of illumination changes. Thus,
central visual processing may proceed without the
brain having to attend to changes in the average
light level caused by the daily solar cycle, by
shading, by artificial illumination, or by other,
perhaps unpredictable, events.
We will demonstrate the visual significance of the
retinal regulation of contrast sensitivity at different
levels of illumination. This basic function of retinal
adaptation is so important for vision that there is
a hierarchy of retinal adaptation mechanisms at
several different sites within the retinal network.
There is clear evidence for adaptation in individual
photoreceptors, in some species. There is evidence
for adaptation at the level of the outer plexiform
layer of the retina, in bipolar cells. There is also
evidence for another stage of adaptation at the inner
plexiform layer, in amacrine cells. Adaptation
performed by the retinal network thus appears to
involve at least three mechanisms in most retinas.
Such evidence leads to the concept of a hierarchy
of mechanisms which may be engaged at different
background levels and with different time courses.
In individual cells or in the retinal network, the
neural signals sent on to the next stage in neural
processing usually are fed back to regulate the
response to new or persistent inputs. The theories
for adaptation which we shall discuss require
feedback in order to explain the phenomena
associated with visual adaptation. One can therefore
state as an overview that visual adaptation is
achieved by a hierarchy of feedbacks designed to
regulate contrast sensitivity.
It is impossible within the space of a review
chapter to deal with visual adaptation to changes
in illumination and also to do justice to the subject
of recovery of sensitivity in the dark after all light
has been turned off. The latter phenomenon usually
goes by the name dark adaptation. Dark adaptation
is in some ways similar to light adaptation but is
different in such significant other ways that it
deserves a chapter all its own. It is not covered in
the following pages.
1.1. Terminology
Because the facts and theories of visual
adaptation are complicated enough, one ought to
be clear about the meanings of words which are
used to describe the facts, and so we will define
several words which are critical for the ensuing
discussion. It is most important to define what
adaptation means, but some preliminary terms
require definition first.
1.1.1. SENSITIVITY AND GAIN
Unfortunately, "sensitivity" has different
meanings in different fields. In psychophysics it
means l/threshold or, in other words, the reciprocal
of the stimulus strength required for the stimulus
to be perceived reliably. According to this meaning,
"sensitivity" is related to the signal/noise ratio
inside the psychophysical observer (cf. for example,
Barlow and Levick, 1969; or Rose, 1948, 1973). The
"noise" in this case is caused by all the physiological
fluctuations in the retina and brain, fluctuations
which make it difficult for an observer to be certain
that a stimulus has been presented. This "noise"
is caused, in the dark, by thermal breakdown of
photopigment in photoreceptors, spontaneous
r a n d o m release o f n e u r o t r a n s m i t t e r s , and
fluctuations in the physiological state of the retina
and brain. When the retina is illuminated,
additional noise is caused by the retinal response
to the randomly arriving stream of light quanta.
Psychophysical "sensitivity" can be influenced by