VISUAL ADAPTATION AND RETINAL GAIN CONTROLS 279

-5 I I I I

-4
: 5 ° Para'[-oveo
;% = 580 nm
# = 5 0 0 nm

-5

-6
,~..
IIII/
o/
I
I
I
--8 1 I I I I
(Zero) -9 --8 --7 --6 --~ --4 --3 --2 -I

Log M,~

FIG. 8. The rod cone transition in h u m a n vision. The log of the energy per unit time per unit area of a test spot is plotted
-

on the ordinate. The abscissa is the log energy per unit time per unit area of a concentric background disk. The units for
both are ergs (deg 2 s)-t measured at the cornea (before losses in the eye and retina). These can be expressed equivalently
as quanta(507 nm) (deg 2 s)-I a n d therefore compared to retinal illuminations in trolands, td (cf. Table 1). The r o d - cone
break takes place at about - 6.7 log ergs (deg 2 s) -1 which is equal to 6.10" quanta(507 nm) (deg 2 s)-1, and approximately
equivalent to 0.1 td. In this experiment the test stimulus was a 1 deg spot presented for 60 ms. The test wavelength was
580 nm. The background wavelength was 500 nm, presented continuously. Background size was not given but was probably
about 10 deg in diameter, concentric with the test spot. Test and background spots were placed 5 deg from the fovea. F r o m
Wyszecki and Stiles (1967).

compared to the variance which is internal to the
retina caused by thermal isomerization o f
photopigment, spontaneous opening and closing of
photoreceptor membrane channels, or spontaneous
neurotransmitter release. These sources of noise
have been shown to act in a way comparable to
illumination, summing over distance and time, and
therefore are conceived of as an equivalent " d a r k
light" (Barlow, 1957).
The value of " d a r k light" was estimated by
Barlow (1957) as the retinal illumination at the
intersection o f a line drawn through the initial flat
portion of the i n c r e m e n t - t h r e s h o l d curve with a
line extrapolated down from the straight sloping
portion (on l o g - l o g coordinates). Let us make
explicit why this intersection gives an estimate of
the " d a r k light". The i n c r e m e n t - t h r e s h o l d curve
PRR3-H
may be expressed in an equation: MT = (ID + IB)p.
P is the slope of the rising portion of the curve on
l o g - log coordinates, and varies from 1 for Weber's
Law to 0.5 for the square root law. The initial flat
portion of the curve is where Io is much larger
than IB and therefore in this range A I x = (Io) P. The
straight sloping portion is where IB is much larger
than Its, and there M x = (IB)P. At the intersection
of the two lines (on log - log coordinates), the two
expressions for the threshold illumination are equal,
and thus at this point the background illumination
Ia equals the " d a r k light" Io.
It appears that the " d a r k light" limits visual
sensitivity by being the dominant term in the retinal
noise rather than by setting the value of the retinal
gain control. Barlow (1957) surveyed the literature
on measurements of " d a r k light" and found that