Dark Adaptation.: R - M - Shapley and C. Enroth-Cugell

264 R.M. SHAPLEY AND C.
ENROTH-CUGELL
1. INTRODUCTION feedback in order to explain the phenomena

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

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264 R.M. SHAPLEY AND C.

1. INTRODUCTION feedback in order to explain the phenomena

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