298 R. M . SHAPLEY AND C.

ENROTH-CUGELL

I I I I I i I I I I I _I

2.5x10 s 0.0~4 2"5 x 105 0.0004

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o"
o v o
~ 2.5x 104 0.~4 2 5 x 10" 0.004

E 2"5 x 103 0-04

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2.5 x 102 i , 0.04
m

I I I I 1
102 104 10' 10 = 104 10'
Backgr. ret. ilium, q(507 nm)/sec, deg 2 Backgr. rec. ilium, q(507 nm)/sec, deg 2

FIG. 24. Gain vs background illumination, in the scotopic range, for cat retinal ganglion cell centers. Retinal flux [in units
of quanta(507 nm) s-'] required for a small central stimulus to elicit a criterion response (30 impulses s -~) is plotted vs the
retinal illumination of a 12 deg concentrically located background. The gain in impulses/quantum (i/q) is indicated also
on the right hand vertical scales. The latter quantity was calculated by multiplying the stimulus retinal illumination by the
area of the stimulus, 0.03 deg 2, and by a factor of 1/3, the estimated fraction of quanta incident on the retina which were
absorbed. In the left hand panel, the filled circles are for white stimuli on a white background; the open circles are for
b l u e - green stimuli on a red background, to demonstrate rod isolation. In the right panel all the points are for white on
white. The results in the left panel are from an on-center ganglion cell; the results in the right panel are from an off-center
cell. From Enroth-Cugell and Shapley (1973a).

because both the " d a r k light" and the transition
illumination are needed to account for observed
plateaus: the background illumination must exceed
IRO for the gain to drop from its dark adapted
value in ganglion cells, while the background
illumination must exceed I D, the " d a r k light", for
the psychophysical sensitivity to drop from its dark
adapted value. However, the functional difference
outweighs the apparent similarity. The transition
illumination is involved in gain control; the " d a r k
light" (either estimated from psychophysics or from
physiological experiments) sets the noise level of the
retina in the dark. This argument is supported by
the estimated values of " d a r k light" and the
transition illumination, which are quite different.
The "dark light" of cat ganglion cells was estimated
by Barlow et al. (1971), as follows. Based on the
value of the maintained discharge in the dark, and
the slope of the stimulus - response curve obtained
in the dark, these authors estimated the magnitude
of the light flux which would have been required
to generate the maintained discharge in the dark,
and called this value the " d a r k light". They found
a wide variation in this estimate of the "dark light".
However, taking their highest value, the " d a r k
light" was equivalent to about 100 quanta s-1 retinal
flux. For a cell with a small center, say about
0.1 deg 2 in area, this would be produced by 103
quanta(deg 2 s)-t retinal illumination; for the largest
cells it would be produced by about 3 q(deg 2 s) -1.
These values for the feline "dark light" are too low,
by at least a factor of ten, for the " d a r k light" to
be equivalent to the transition illumination in cat
retinal ganglion cells. Rather, some criterion
amount of voltage or current or substance in a
retinal cell, much larger than that caused by " d a r k
light", must be exceeded, and then the gain control
of adaptation begins to act. As argued in Section
2.1.1.2., "dark light" probably limits sensitivity by
providing a noise, the "dark noise", against which
a signal must be picked out, rather than by setting
the gain.
3.1.2. GAIN AND DYNAMICS
There are dynamic consequences of adaptation
which are hidden in the simple picture of Fig. 24.
As shown in Fig. 25, the time course of response
of the receptive field center varies with adaptation
level, as found both by Yoon (1972) and by Enroth-
Cugell and Shapley (1973a). The nature of the
change is that the response of the center to an
incremental step of illumination becomes more
transient, the more light adapted the cell is in the