328 R. M. SHAPLEY AND C.

ENROTH-CUGELL

r e s p o n s e to a d a p t a t i o n ( T r a n c h i n a , G o r d o n and (1974). F i g u r e 56 s h o w s t h e i r results. It c a n be seen

Shapley, unpublished). t h a t the r o d o p e r a t i n g curves shift to the right a very
msec little w i t h i n c r e a s i n g b a c k g r o u n d , b u t m a i n l y
0 1 O0 200 3 O0 collapse due to saturation. Similar receptor
I I I b e h a v i o r is i m p l i e d b y t h e h o r i z o n t a l cell results o f
30 -
W i t k o v s k y (1967) in c a r p a n d N a k a et al. (1979) in
c a t f i s h . R e c e n t l y , N u n n a n d B a y l o r (1982) h a v e
r e p o r t e d t h a t r o d s in t h e m o n k e y Macaca
10 fascicularis m a i n l y s a t u r a t e w i t h o u t a d a p t i n g . T h e r e
is s u p p o r t i v e e v i d e n c e f r o m mass r e c e p t o r p o t e n t i a l
0 4"52 ~ ~ " " "; r e c o r d i n g in t h e rat t h a t at least s o m e m a m m a l i a n
rods saturate without adapting (Penn and Hagins,
1972; G r e e n , 1973). T h i s is also c o n s i s t e n t w i t h the
v e r y s m a l l a m o u n t o f a d a p t a t i o n in t h e a - w a v e o f
2
I- the r a t ' s E R G , t h o u g h t to be d e t e r m i n e d by r e c e p t o r
E a n d h o r i z o n t a l cell r e s p o n s e s ( D o w l i n g , 1967), a n d
o 1
b y m a s s r e c e p t o r p o t e n t i a l r e c o r d i n g in t h e cat eye
0
~. 0 -~ in w h i c h the i n n e r n u c l e a r layer a n d t h e g a n g l i o n
> cell layer w e r e r e n d e r e d a n o x i c by o c c l u d i n g t h e
5.79 o p h t h a l m i c a r t e r y ( S a k m a n n a n d F i l i o n , 1972).
0.15 S a k m a n n a n d F i l i o n also s h o w e d t h a t t h e g a i n o f
the isolated rod receptor potential was

0-10

0.05
E=2

Q ~
0 - I ,,I ' :

1 ~ I
I oo 200 300
msec

FIG. 55. Effect of increasing illumination of conditioning
step on the time course of the flash response in a turtle cone
(intracellular), The flash was I1 ms in duration, and was
applied 1.1 s after the onset of a conditioning step of
illumination. The horizontal axis is the time after flash onset.
The vertical axes are all in units of gain:
/aV/photoisomerization, which is equivalent to taV/quantum
effectively absorbed. The numbers near each curve are the
illuminations of the conditioning steps, in units of
log[effective quanta(cone s)-q. These data are from a red
cone. The stimulus was a 150/am white spot. From Baylor
and Hodgkin (1974).
5.2. Photoreceptors which Saturate but do not
Adapt
T h e best s t u d i e d e x a m p l e s o f r e c e p t o r s w h i c h
s a t u r a t e w i t h o u t a d a p t a t i o n a r e the r o d s o f t h e
mudpuppy studied by Normann and Werblin
04
Fro. 56. Rod operating curves on backgrounds. Plotted is
the magnitude of the rod response (recorded intracellularly)
measured as the difference between the peak of the response
and the steady polarization just before the response. The
response to test flashes brighter than the background are
shown below the horizontal axis, and to test flashes dimmer
than the background above the axis. The stimuli were diffuse
2 s flashes. The background illuminations can be read as the
test stimuli which produced zero response, i.e. the intersection
of the curve with the horizontal axis. Circles are from a rod
in a normal retina, and squares are from a rod in a retina
treated with aspartate. The units of the horizontal axis, which
indicates the test flash illumination, are as follows: 6. l0 s
quanta(522 nm) (cm2 s)-1 when log I = 0. From Normann and
Werblin (1974).