VISUAL ADAPTATION AND RETINAL GAIN CONTROLS 299

qQ t/Q
ratio -J.e--1.25 sec ~l [ On ratio

---IOn I I~

0.08

0-01
002. I50 impulses/see

0006 150 impulses]sec

(a) (b)

FIG. 25. Time course o f (rod-driven) square wave responses at different adaptation levels in cat retinal ganglion cells.
(a) The responses (averaged over many stimulus cycles and smoothed) of the receptive field center of an on-center cell at
five different levels of background retinal illumination: (a) l - - 5 0 quanta(deg 2 s) "1, (a)2--6. l & quanta(deg a s)-', (a)--6" l&
quanta(deg 2 s) -1, (a)4---9.104 quanta(deg 2 s)-t, and (a)5--5" l& quanta (deg 2 s) -1.
(b) A similar result is shown for an off-centercell at three backgroundilluminations: (b)l--100 quanta (deg2 s)-t, (b)2--3. l&
quanta(deg2 s)-l, (b)3--103 quanta (deg2 s)-'. Note that in the cat 1 deg2=0.048 mma, approximately.
For each response the i/q ratio, the gain, is shown at the left of the averaged response histogram. The time courses of
the responses and the i/q ratios at the top of each column were the same as those obtained in total dark adaptation. The
drop in gain with increase in background illumination goes hand in hand with the change in time course observed in the
histograms. From Enroth-Cugell and Shapley (1973a).

scotopic range. This finding applies to off-center
as well as to on-center cells, and to X as well as Y
cells (Jakiela et al., 1976).
This change in time course of response with
adaptation is related to another observation: the
steepness of the decline in gain with background
depends on the temporal pattern of the stimulus
(and response). For example, the results in Fig. 24
were derived from measurements of the peak
response to an incremental step of illumination on
a background. One would obtain rather similar data
from measurements of the gain o f the response to
a sinusoidally modulated small spot for temporal
frequencies of 2 Hz or less. In this case, either peak-
to-peak impulse rate modulation or the amplitude
of the sinusoid which is the best approximation to
the neural response are two response measures
which would give the same dependence on
background. Similar measurements at 8 Hz or
above have a shallower dependence on background,
i.e. the exponent P in equation (20) would be
around 0.6 for intermediate temporal frequencies
of modulation (between 3 and 10 Hz; Enroth-
Cugell and Shapley, 1973a; Derrington and Lennie,
1982). The responses to temporal frequencies above
16 Hz suffer almost no attenuation in amplitude
with increases in mean level; for such high temporal
frequencies, the exponent P in equation (20) is near
zero (Shapley et al., 1983). Previously, Sakmann
and Creutzfeldt (1969) and Barlow and Levick (1976)
also observed shallow slopes of gain vs background
curves with brief incremental stimuli. The responses
to such brief pulses of light, which contain a wide
range of temporal frequency components, follow
a gain vs background curve which obeys equation
(20) with an exponent of about 0.6 on the average.
However, as presented earlier, the gain of the peak
of the response to a prolonged flash has a steeper
d r o p o f f of gain with background; the exponent in
this case is about 0.8, according to Barlow and