VISUAL ADAPTATION AND RETINAL GAIN CONTROLS 325

5.1. Photoreceptors which Adapt a lot and Saturate

The photoreceptors which actually adapt, i.e.

more or less rapidly adjust their gain to avoid 10
saturation, include the following rods: skate,
s
(Dowling and Ripps, 1972); gecko, (Kleinschmidt
and Dowling, 1975); toad (Fain, 1976; Baylor et al., o
o.
1979). The adapting cones include: turtle (Baylor
and Hodgkin, 1974; Normann and Perlman,
1979a); mudpuppy (Normann and Werblin, 1974); 1
frog (Hood and Hock, 1975); ground squirrel
(Dawis and Purple, 1982); perhaps monkey O5
I I I I I I
(Valeton and van Norren, 1983). The results on 7 8 10 11 12 13
cone adaptation in ground squirrel, frog, and log (quanta crn -2 flash 1)

monkey are from massed potential recordings which
in one way or another isolated the cones.
Figure 50 illustrates photoreceptor adaptation in
rods from the toad B u f o marinus (Fain, 1976).
There is evidence of saturation (in the decline of the
peak response on background illumination), but
there is also clear curve-shifting of the V-log/curve,
consistent with the idea of an automatic gain
FIG. 50. Effect of background light on the intensity-
response curves of toad rods; intracellular recording. Peak
response amplitude to a brief (around 100 ms) flash is plotted
against log flash energy per unit area (quanta cm-2), for the
dark adapted rod and for several background levels of
illumination. The stimulus and background were diffuse. The
illumination of the backgrounds is given in log units to the
left of each curve, and the units are log quanta(505 nm)
(cm2 s)-1. From Fain (1976).
100 - -

control, equation (10) (cf. Normann and Werblin,

1974; Dawis and Purple, 1982). Furthermore, the
gain of the toad rod follows Weber's Law (Fain, lo ~ s d

1976), and this can even be seen in the photocurrent

recorded from the outer segment by the suction
electrode technique (Baylor et al., 1980) as in '~
Fig. 51. In Fig. 51 the vertical axis is labeled ~ lo
"sensitivity", but it is equivalent to what we have _
defined as "gain". Log gain vs log background has
a flat portion and then a declining portion with a 01
slope about - 1. The slope of unity suggests an
adaptive gain control rather than the steeper slope
associated with pure saturation (see Section 1.2.2.). 7"." I ! I I I
The transition from flat to sloping is at about an 0-01 0"I I0 10 I(}(3
le (photons ~ m -2 sec -I)
illumination of 0.28 quanta/am-2 s-1, which is about
forty times greater than the "dark light" of the FIG. 51. Gain vs background in the photocurrent of a toad
rod outer segment; recorded with suction electrode. The gain
rods, the spontaneous current fluctuations in the is plotted on the vertical axis with units picoamps/(photon
dark, measured independently. Baylor et al. (1980) /~m-2), but could really be given in picoamps/photon since
make the point that this implies that the gain control the rods are of fixed cross-sectional area. The background
illumination is plotted on the horizontal axis. The transition
transition level in rods is not determined by the from horizontal to sloping portions of the curve occurs at
"dark light". Similarly, we have pointed out in 0.28 quanta(507 nm) (/am2 s)-1. The arrow points to this
connection with gain reduction in ganglion cells, illumination. Background and test were diffuse. From Baylor
et al. (1980).
that the "dark light" in the cat retina is too small
to be responsible for determining the illumination the rod response. The gain declines by about a
at which gain starts to fall (see Section 3.1.1.). factor of one hundred over a range of backgrounds
The gain reduction of the rod photocurrent is which speed up the time to peak of the photocurrent
accompanied by a very significant speeding up of (in response to a brief flash) by a factor of seven.