VISUAL ADAPTATION AND RETINAL GAIN CONTROLS 277

NERISSA
When the moon shone we did not see the candle.
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PORTIA
So doth the greater glory dim the less.
A substitute shines brightly as a king
Until a king be by, and then his state
Empties itself, as doth an inland brook 0 -- 4
Into the main of waters . . . . g
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(Act V, Scene 1, II. 89-97).

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T h e basic p h e n o m e n o n is a c c u r a t e l y d e s c r i b e d 4-"
here, the f a d i n g to d i s a p p e a r a n c e o f a w e a k
E
I1) - I -
i n c r e m e n t o n a b a c k g r o u n d . Since the b a c k g r o u n d
b B
was m o o n l i g h t in S h a k e s p e a r e ' s " e x p e r i m e n t " ,
N e r i s s a m u s t have b e e n o b s e r v i n g light a d a p t a t i o n
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o f the r o d system. This we n o w c o n s i d e r m o r e
quantitatively.
Scare 06
-31
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2.1. The Rod System
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2.1.1. SENSITIVITYAS A FUNCTION OF BACKGROUND Log ( f i e L d radiance,scotopic ±roLands)

2.1.1.1. Aguilar and Stiles' experiment. A basic

fact o f visual experience is t h a t the sensitivity for
increments o f light is reduced b y steady b a c k g r o u n d
i l l u m i n a t i o n . W e will review the p s y c h o p h y s i c a l
investigations o f this p h e n o m e n o n in o r d e r to
a n s w e r the q u e s t i o n , " t o w h a t extent is the loss o f
visual sensitivity during i l l u m i n a t i o n caused by true
light a d a p t a t i o n o f the visual system, a n d to w h a t
extent is it caused b y i n c r e a s e d " n o i s e " in the
stimulus i t s e l f ? "
The canonical d a t a concerning the d e p e n d e n c e o f
the p s y c h o p h y s i c a l t h r e s h o l d o n the level o f
b a c k g r o u n d i l l u m i n a t i o n are t h o s e o f A g u i l a r a n d
Stiles (1954; Fig. 7). T h e stimuli were c h o s e n to
s t i m u l a t e the r o d system m o s t effectively. T h e
stimulus was large (9 ° d i a m e t e r disk), p r e s e n t e d in
the p e r i p h e r y o f the visual field, a n d was o f a
b l u e - g r e e n c o l o r o n a red b a c k g r o u n d . T h e red
b a c k g r o u n d desensitized the cone system relatively
m o r e t h a n w o u l d a white b a c k g r o u n d , a n d the
b l u e - g r e e n test d i s k s t i m u l a t e d the r o d s m o r e
effectively c o m p a r e d to cones t h a n w o u l d a white
stimulus. T h e stimulus was p r e s e n t e d for 0.2 s
which a l l o w e d the r o d s to s u m m a t e their signals
temporally. In this way, A g u i l a r a n d Stiles were able
FIG. 7. Human " r o d " threshold as a function of
background illumination. The stimulus was a 9 deg disk on
a 20 deg background presented 9 deg from the fovea. The
disk was blue - green and the background was red in color,
in order to enhance rod responses compared to cone
responses. The stimulus was presented for 200 ms. The
background was continuous. Curve A is the threshold
illumination vs background. Curve B is what we would call
the Weber contrast at threshold, and is labeled the "Fechner
fraction" in this graph. From Wyszecki and Stiles (1967);
original data replotted from Aguilar and Stiles (1954).
to isolate the r o d system a n d s t u d y its p r o p e r t i e s
over a wider range o f b a c k g r o u n d s t h a n has usually
been possible. O n e unit m u s t be d e f i n e d in o r d e r
to u n d e r s t a n d their results, the unit o f retinal
i l l u m i n a t i o n , the t r o l a n d ( a b b r e v i a t e d td). In
h u m a n s , a l u m i n a n c e o f 1 cd m -2 viewed t h r o u g h
a p u p i l with a n a r e a o f 1 m m 2 is said to g e n e r a t e
a retinal illumination o f 1 td. The values o f trolands
in m o r e p h y s i c a l units is given in g r e a t detail b y
W y s z e c k i a n d Stiles (1967).
A s A g u i l a r a n d Stiles d e s c r i b e d their d a t a , the
first m a j o r f e a t u r e s h a r e d b y several observers was
t h a t w h e t h e r t h e r e was zero b a c k g r o u n d o r a
b a c k g r o u n d b e l o w 2 . 1 0 -3 t d , t h e t h r e s h o l d