326 R. M. SHAPLEY AND C.

ENROTH-CUGELL

I I I I I I I
• Dark adapted
-15 • -4"4 BG
D -3.2 BG
• -2.1 BG

-10

g -5
g.
• • 121
0 ........................................................................

I I I I I I I
--6-0 --5"0 -4.0 -3"0 -2.0 -1.0 0.0
Log relative intensity

FIG. 52. Gain adjustment in turtle cones (recorded intracellularly) caused by background illumination. The stimuli were
0.5 s increments or decrements on a steady background (except for the curve for the dark adapted cone which only is for
increments). The stimulus spot was 3.2 mm in diameter on the retina. Peak responses measured from the dark-adapted
resting potential (dotted line) were plotted as a function of test illumination. The curves are Naka- Rushton functions.
The horizontalbar through each curveindicatesthe steadymembranepotential measuredat least two minutesafter background
onset. The illuminations are given as log attenuation. The unattenuated test stimulus (0 log) was 6.4.10" quanta(640 nm)
(cm2 s)-' on the retina. The unattenuated background illumination was 9.1.10 is quanta(640 nm) (cm2 s)-L From Normann
and Perlman (1979a).
Baylor e t al. (1980) f o u n d that G R ~ (tpeak)25, 2-
where G R is the g a i n o f the r o d in
p i c o a m p / q u a n t u m a n d tpeak is the time from the / ~ 0
onset of the brief flash stimulus to the peak o f the 1

response.
A d j u s t m e n t of gain in cones by a d a p t a t i o n a n d
s a t u r a t i o n is represented in Fig. 52 from the data
of N o r m a n n a n d P e r l m a n (1979a). The parallel ~ 0
curves are the response template f r o m the dark ,-I
a d a p t e d cone, a n d are described by the N a k a -
R u s h t o n r e l a t i o n , e q u a t i o n (6). W h e n the
b a c k g r o u n d is raised, the response to increments is
somewhat compressed (consistent with s a t u r a t i o n )
because the steady state response increases towards
the m a x i m a l potential the cone is capable o f
producing, as indicated by the short horizontal bars
intersecting each operating curve. F r o m the shift o f
the operating curves, one m a y infer W e b e r ' s Law
since there seems to be a b o u t a one log unit shift
for each log unit increase in background. That turtle
cones a p p r o x i m a t e l y follow W e b e r ' s Law is
indicated in Fig. 53 from Baylor e t al. (1974b). The
solid line is W e b e r ' s Law u p to a b o u t log Is = 4.5,
a n d t h e n rises due to s a t u r a t i o n . It is based o n a
theory for the c o n e ' s gain c o n t r o l m e c h a n i s m by
Baylor e t al. (1974b). A c t u a l l y the c o n e ' s gain
follows W e b e r ' s Law for a b o u t a log unit more t h a n
the Baylor e t al. theory predicts, a n d saturates
rather less t h a n the theory predicts. One possibility
n o t considered by Baylor et al. is that some o f the
/
-
I
_o
-2
-3 I I I I I
2 3 4 ,5 6 7
log I,
FIG. 53. Turtle cones' gain as a function of background;
intracellular recording. The ordinate of this unique plot is
the logarithm of the difference between the reciprocal of the
gain in the light adapted cone and the reciprocal of the gain
in the dark adapted cone, i.e. Iog{1/SF-I/SFD}. This number
unfortunately goes to minus infinity when the background
is so low that the gain of the cone is the same as in the dark
adapted state, but that region of the curve is not plotted here
anyway. The gain SF is in gV quantum-' effectively
absorbed. The horizontal axis is log background flux; the
units are quanta/sec effectively absorbed by the cone. The
stimuli were 10 ms flashes, and the gain is calculated for the
peak of the flash response. These are data from a red turtle
cone. The smooth curve is from Baylor et al.'s theory for
cone adaptation. From Baylor et aL (1974).