VISUAL ADAPTATION AND RETINAL GAIN CONTROLS 327

deviation at the high backgrounds might be due to
pigment bleaching, which is not included in their
theory. Cones also speed up their response with
background and for turtle cones, Gc ~" (tpeak)4, that
is the gain of a cone decreased like the fourth power
of its time to peak for the response to a brief flash.
The total amount of "speeding up" of the cone
response is about a factor of two over a range of
backgrounds in which the gain drops by a factor
of two hundred. The speeding up of rod and cone
responses with the change in gain is illustrated in
Fig. 54 (Baylor and Hodgkin, 1974; Baylor et al.,
1980).
Steady illumination affects different parts of the
cone's response in different ways. This is illustrated
in Fig. 55 from Baylor and Hodgkin (1974). The
I I
impulse response of the cone, i.e. the response to
a brief, weak flash of light in the linear range,
changes as the steady background increases. The
gain for later parts of the impulse response drops,
but the gain for the initial rising phase of the
response is relatively unaffected by steady
backgrounds. The peak of the response becomes
smaller at higher backgrounds and occurs at earlier
times after the flash, but the rising phases of
impulse responses at several levels of background
superimpose. Although this feature may not be
completely convincing in Fig. 55, it is true. Recent
preliminary experimental results involving the
measurements of temporal frequency responses in
turtle horizontal cells and cones are consistent with
the resistance of the rising phase of the impulse

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100 -

50-

20-
• O
10-
E

0 E 5.0-
O
O
f~
2.0-
_o
• -
1.0
--1
0-5

0.2

--2 0.1 I I I I I
0-1 0-2 0.5 1.0 2.0 5.0 10
f'a~ak (see)

I , I I
1.6 1.8 2.0 2.2
log tmax
FIG. 54. The speeding up of cone and rod responses in the turtle and toad retinas. As mean illumination increases, the
gain and the time to peak decrease. In the left panel, the log of the gain is plotted against the log of the time between
the peak response to a brief flash and the onset of the flash, for turtle cones (intracellular recording). The gain is plotted
against peak time for toad rods (suction electrode recording) in the right panel. For about two log units log tmax is
approximately proportional to log gain (here denoted SF). In the left panel, the different symbols are for different red
cones, eight in all. On the right are graphed data points from seven different rod outer segments. Note that the cone time
to peak stops decreasing when the gain drops below 0 log. The units of the left ordinate are log[#V (quanta gm-~)-l]; on
the right the units are picoamps (quanta ~m-2)-L On the left the units of the abscissa are log[ms]; on the right they are
simply s. From (left) Baylor and Hodgkin (1974), and (right) Baylor et al. (1980).