VISUAL ADAPTATION AND RETINAL GAIN CONTROLS
z,,(co) y.l(co)
I,(o~) I,(co) (33
1 Hodgkin (1974). They found that the rising phase
+ A
1 + io~B2
where T = l / a
B, = [g,2 + K2,]/[K23"K,2]
Bz = 1/{J~2,[1 + vlo/(Kz3K,2)]}
A = K,2/K2,.
Notice that the frequency response of the cone
consists of three terms, the first two of which are
stages of temporal integration which are indepen-
dent of the mean level, Io. The only place where io
enters in the expression for the cone's light adapted
frequency response is in the time constant B~ in third
term of equation (33). This term has the form of
the frequency response of a negative feedback loop
which has a gain A and time constant B2. The time
constant of the feedback is what is affected by mean
level in the BHL model. As Baylor et al. (1974b)
demonstrated, this could account qualitatively for
the effect of backgrounds on the gain and time
course of the cone's impulse response [the Fourier
transform of the frequency response in equation
(33)]. However, as indicated above in Fig. 53, the
predicted gain vs background curve is steeper than
the real data, while the dependence of the time to
peak of the response on mean level is too shallow.
In a later paper, concerning toad rods, Baylor et
al. (1980) proposed that feedback to earlier stages
of processing, rather than to just the last stage as
in the BHL model, would provide a better fit to the
data. However, the theory of such a system has not
been analyzed and would certainly be more
complicated than the BHL model.
A very important consequence of the BHL model
is that the amplitudes of responses to high temporal
frequencies of modulation are unaffected by mean
level. This can be seen in equation (33) in the limit
as co~oo. Then the only term which contains Io
approaches the value unity, and it does so at lower
values of co than the other terms approach zero.
333
Thus, the BHL model predicts no effect of light
adaptation in the limit of high frequency. This is
qualitatively in agreement with the measurements
of the cone's impulse response by Baylor and
of the impulse response was not affected by
background level. This result is also qualitatively
in agreement with Kelly's (1972) psychophysical
data on the linearity of high frequency response in
h u m a n vision. R e c e n t direct i n t r a c e l l u l a r
measurements of the frequency responses of turtle
horizontal cells also confirm this prediction of the
B H L theory (Tranchina, Gordon and Shapley
unpublished).
6.1.3. ENROTH-CUGELL AND SHAPLEY'S MODEL OF
ADAPTATION IN GANGLION CELLS
Another scheme for obtaining Weber's Law
behavior linked to dynamic changes is diagrammed
in Fig. 60 (Enroth-Cugell and Shapley, 1973a). In
this model for the rod-driven retinal network of the
cat, the rod signal is subjected to a multiplicative
gain control at the level of the r o d - b i p o l a r
synapse. In the original model it was proposed that
the feedback signal which multiplied the rod signal
was produced by the horizontal cell. Later work
reviewed above indicates rather that the feedback
signal may arise in bipolar cells. In any case the
formal expression of the model is:
r(t) = P(t)/exp{H(t)/Htrig}
P(t) = f ~ ( t - t ' ) p ( t ' ) dt'
p ( t ' ) = g - ( t ' / r , ) 3 exp { - t ' / x p } (34)
H(t) = f ~ r ( t - t ' ) h (t') dt'
h ( t ' ) = exp (--t'/TH).
Where r(t) is the rod signal to the bipolars, H(t)
is the horizontal cell potential (now thought of as
the bipolar cell's potential) and T. is the time
constant of the feedback neuron. Htri, is a critical
level this signal must exceed, Tp is the time constant
of the rod, I(t) is the light stimulus, P(t) is the
photocurrent of the rod, ~ is the gain in the dark.
The frequency response of this model has been
worked out and it is (Enroth-Cugell and Shapley,
1973a):