334 R.M. SHAPLEY AND C.

ENROTH-CUGELL

r,(¢o) (35)
- g-exp {-ro/Htrig} n

:E pj(t)
l,((o) j=o

f l +T io)t 4 ~ + 1l ' ° n t rig t

1 + im'rn.~

where ro is the steady state value of r(t). In the

model, ro approximately increases like log Io for
g-Io>>H~r~g. Two features are noteworthy. The
general " s h a p e " of the frequency response
resembles that of the BHL model's frequency
response, equation (33). However, the effect of n ell(t)
(t)=~rj(l)
mean level on dynamics is on the gain of the
feedback term ro Htrig, rather than on the time I
ex
constant r,,. Moreover, the gain of high frequency
responses in this model does drop at high
backgrounds by the factor e x p { - r o / H , ng}, that is
approximately reciprocal to Io in the high
background limit. This prediction is not correct for (tl ~"-I
single cone responses, but it has not been tested in Hot celt J
the system for which this model was designed, the r(~;; r
cat retina in the scotopic range. Measurement of
the temporal frequency response of cat retinal
ganglion cells at several different background or
FIG. 60. A model for adaptation in the cat retina. The p(t)s
mean levels would be a crucial test of the different are the photocurrentsin individualrods. The amount of signal
predictions of the BHL model and the E n r o t h - transmission from rods to bipolarand horizontalcellsis called
Cugell - Shapley model. Preliminary experiments of r(t), with subscripts for each rod. H(t) is the horizontal cell
potentialdue to the sum of its inputs. An exponentialfunction
this type appear to confirm a model of the BHL of the horizontalcell'spotentialis what controlsthe sensitivity
type and to exclude multiplicative models like the of receptor transmission. Thus, r(t) =p(t )/exp{H(t )}. From
E n r o t h - C u g e l l - Shapley model (Shapley et al., Enroth-Cugell and Shapley (1973a).
1983).

6.1.4. THE CONTRAST GAINCONTROL MODEL

The contrast gaincontrol adjusts the time course
of response contingent on the average level of
contrast rather than simply contingent on the mean
flux. Shapley and Victor (1981) proposed a theory
r~(o))

11(¢o)
(36)
-
A

f 1

+ i¢o

Shapley and Victor (1981) found that increase of

+

contrast affected only the ratio K/tH and had only

for the way in which the time course of response
was changed by contrast, a theory which resembles
the BHL model. Their model is diagrammed in Fig.
61. There are n stages of temporal integration and
one negative feedback stage. The temporal fre-
quency response of this system is:
1 +- iwT
1
negligible affects on A, x~ or n. Formally, equation
(36) resembles equation (33) for the BHL model.
The effect of contrast is exerted on the same term,
the feedback term, and in a roughly similar way.
Whether this similarity of the consequences of these
models for incremental responses has any deep