VISUAL ADAPTATIONAND RETINALGAIN CONTROLS
TABLE I. Quantal Equivalents of Photometric Units and some Other Useful
Equivalences
Photometric Unit
1 cd m -2
through 1 mm ~
pupil, scotopic
(1 scotopic td, h u m a n )
1 cd m -2
through 1 mm ~
pupil, p h o t o p i c
(1 p h o t o p i c td, h u m a n )
1 l u m e n (scotopic)
1 lumen (photopic)
Other Useful Equivalences
Unit
1 q u a n t u m ( 5 0 7 nm) s-'
1 q u a n t u m ( 5 6 0 n m ) s -I
1 deg 2 o n h u m a n r e t i n a
1 deg 2 cat r e t i n a
is based on more recent estimates of quantum
efficiency by Barlow, 1977). However, Sakitt (1972)
has shown that if one relaxes the stringent
requirement of no false positives, human observers
can do better than chance when on the average only
a single quantum of light excites the retina. This
must mean that a single quantum response is
comparable in magnitude to the intra-retinal dark
noise. Comparable sensitivity is possessed by other
animals.
The high sensitivity of a dark adapted eye poses
a problem when the observer moves into brighter
surroundings. Almost all neurons have a limited
response range, from small signals to the peak levels
set by biological constraints such as ionic
equilibrium potentials. The range of responses is no
greater than a factor of one hundred from noise to
ceiling. The problem is obvious. How can the rod
pathway in the retina encode three to five log units
of stimulus level when it only has a factor of one
hundred in response to work with? The answer is
it cannot, and it does not. The retina adapts
(reduces its gain) in the presence of large average
inputs in order to represent only modulations
around the average level when the average becomes
too large to handle with the high dark adapted gain.
This problem of saturation of neural responses
is related to the need for stable contrast sensitivity,
which we have argued is the main purpose of
adaptation. If the retina did not adapt, the contrast
gain and contrast sensitivity would plummet at high
273
E q u i v a l e n t in Q u a n t a
4.46. l 0 s q u a n t a ( 5 0 7 nm) (deg 2 s) -t
1.26.106 q u a n t a ( 5 6 0 nm) (deg 2 s) -1
1.4.1015 q u a n t a ( 5 0 7 nm) s -1
4 . 2 . 1 0 is q u a n t a ( 5 6 0 nm) s -1
Useful Equivalent
4 . 1 0 -'9 w a t t s
3.5" 10 -'9 w a t t s
8.5" 10 -4 cm 2
4 . 8 - 1 0 -4 cm 2
light levels. Thus we would become blind to
reflecting objects in bright daylight. This would not
be a stable survival strategy, and therefore there is
a biological need for adaptation. In fact, as our
review of the psychophysical and physiological
results will show, the contrast sensitivity and
contrast gain of humans and animals generally
increase as the illumination increases, finally
levelling off to asymptotic values in bright light. At
this point we will demonstrate how the saturation
o f neural response would lead to a decline in
contrast gain. Then we will present a suggestion for
an adaptation mechanism which offers an escape
from the "saturation catastrophe".
First let us consider how the N a k a - R u s h t o n
equation, which approximately describes the
i n t e n s i t y - r e s p o n s e function of distal retinal
n e u r o n s , would lead to a " s a t u r a t i o n -
catastrophe" if the retina did not adapt. This is the
N a k a - Rushton equation:
R = ( I / ( I + Is)) Rmu (6)
where R is the response o f the neuron measured as
the change in membrane potential from its totally
dark adapted level, I is the illumination of the
stimulus. Is is the semi-saturation constant also
equal to the illumination at which R reaches its half-
maximal value. This equation (Naka and Rushton,
1966) is called the Michaelis- Menten equation by
Baylor and colleagues (Baylor and Hodgkin, 1973;