274 R. M. SHAPLEYAND C.

ENROTH-CUGELL

B a y l o r et al., 1974; B a y l o r et al., 1979) after a the a m o u n t o f response per a m o u n t o f c o n t r a s t ,

similar e q u a t i o n which arises in the t h e o r y o f thus:
enzyme kinetics. T h e e q u a t i o n describes a system
which saturates. Figure 4 shows a g r a p h o f R/Rmax Gcon dR/(dI/Ia)
= = dR/dlog/
vs / p l o t t e d on l i n e a r - linear coordinates, a n d it can = ~/(~r/&) (8)
be seen t h a t R is a s a t u r a t i n g f u n c t i o n o f I: a b o v e
the value I s , the light can increase by several a n d to calculate the c o n t r a s t gain for a n e u r o n
o r d e r s o f m a g n i t u d e b u t the response, R, can which obeys the N a k a - R u s h t o n relation we
increase at m o s t b y a f a c t o r o f two. substitute e q u a t i o n (7) into e q u a t i o n (8) to o b t a i n :

Gcon = /sRmax" Ia/(/B +/s) 2

1.0 - "-~ (/sRmax) " 1/([a) as/B>>Is. (9)

0.8 - ~ W h a t this m e a n s is t h a t on a c c o u n t o f s a t u r a t i o n
g 0.6
the W e b e r c o n t r a s t gain w o u l d decline as the
E
or- a m o u n t o f s t e a d y b a c k g r o u n d light increased, in
0.4 inverse p r o p o r t i o n to the average light level. T h u s
the " s a t u r a t i o n - c a t a s t r o p h e " is i m p l i c i t in
0.2 e q u a t i o n (9). F i g u r e 5 illustrates the gain a n d the

0 I I I I
0 200 400 600 800
ILLUMINATION 1.0 ~/--~,N\x -- Goin
--- C o n t r o s f Getn
D 0.8 xx\
FIG. 4. The Naka-Rushton relation. This is a plot of
(R/Rmax)=I/(I+Is), the N a k a - Rushton relation, on a. 0.6 "-.~
linear-linear coordinates. As can be seen, the relation
produces a compressive nonlinear curve; the response
saturates when I > I s. In this example, the semi-saturation ~ 0.4 ~
illumination, I s, was chosen to be 100 in arbitrary units.
~ 0.2

A s a r g u e d by W i l l i a m s a n d G a l e (1977), N a k a et o "7---~1 1 -..

al. (1979), N o r m a n n a n d P e r l m a n (1979c), a n d
Valeton and van Norren (1983),
N a k a - R u s h t o n e q u a t i o n can be used to describe
gain a n d c o n t r a s t gain in e x p e r i m e n t s in which the
stimulus is an i n c r e m e n t o n a b a c k g r o u n d , b y
examining how an incremental response h R depends
on an i n c r e m e n t a l stimulus h / s u p e r i m p o s e d on a
b a c k g r o u n d I B. In t h i s c a s e t h e I in t h e
N a k a - R u s h t o n e q u a t i o n is IB + h i , the response
is R B+ AR, a n d the gain is d R / d I ~- AR/AI, when
h i is small. The gain can be c a l c u l a t e d by
d i f f e r e n t i a t i n g e q u a t i o n (6) ( W i l l i a m s a n d Gale,
1977; N a k a et al., 1979):
dR/dI = [l/(/a+/s)] 2/sRmax
~ { 1/(&2)}. (/sRmax) f o r / a > > / s
T h e W e b e r c o n t r a s t gain Gcon can be d e f i n e d as
o 200 400 600 800
ILLUMINATION
the
FIG. 5. The gain and (Weber) contrast gain of the
Naka- Rushton relation. Gain is drawn as the continuous
curve and contrast gain as the dashed curve. The saturating
nature of the Naka - Rushton relation causes the gain to be
a monotonically decreasing function of illumination [cf.
equation (7)], while the contrast gain has a peak at I = I s
(which was chosen to be 100 as in Fig. 4).
contrast gain (equations 7 and 9) for a neuron which
obeys the N a k a - R u s h t o n relation. I f the retina
w o u l d not a d a p t , the ( W e b e r ) c o n t r a s t gain w o u l d
d r o p as the light level increased.
T h e retina a d a p t s to a v o i d s a t u r a t i o n by having
an " a u t o m a t i c gain c o n t r o l " (Rose, 1948); the gain
o f the retina is reduced after the reception o f enough
(7) light, so that the n e u r a l response u s u a l l y d o e s n ' t
s a t u r a t e in the p h y s i o l o g i c a l r a n g e o f i l l u m i n a t i o n .
W e will have a lot to say a b o u t the m e c h a n i s m o f
this a u t o m a t i c gain c o n t r o l , w h e n we discuss the