VISUAL ADAPTATION AND RETINAL GAIN CONTROLS 331

BuiLd up of particles

Q u (i (~
~(-t) m,,,~l~- y, 1 Y2 D Y3 l l l l l ] l ~ Y4 ]1 Y5 " Y6
Light
(=Z,)

Removal of part-ictes

K1 2 ~ " ~ K23 K34

m ~ m dll~ Z~ Z2 . Z3 I Z4
K21 K32
Open channels

ChanneLs
closed by Z,

FIG. 59. This is a diagram of the Baylor- H o d g k i n - Lamb (BHL) model. The blocking substance is denoted y~ or z,. Y
is used for build-up, z for destruction of the blocking substance. Note that this scheme is fundamentally different from
the FH model (Fig. 58) in that the rate constant for production of each y stage is the same as for its decay, except for
the first and last stages. In the FH model, the rate constant for growth of each stage is different from the rate of decay,
and only the decay rate is subject to change by adaptation. Here adaptation only works by changing the rate constant of
decay from state z,. From Baylor et al. (1974).

where Yis the gain of the initial transduction from
fight to y and I(t) is the fight stimulus. The last stage,
the time evolution of which is given by y,(t), is
presumed in the model to control the membrane
conductance and thereby the electrophysiological
response to light o f the photoreceptor. In
comparing equations (25) and (26) with Fig. 58, one
should make the identification a , = ~ / C and
a2 = 1/RC.
This model was developed to explain first of all
the many stages of temporal integration in the
process of visual transduction, as reflected in the
slow rise to the peak of the impulse response of
photoreceptors. It does this very well. In turtle
cones and toad rods the model indicates that the
number of stages is from four to six. From the
experimental evidence of the toad rod photocurrent
recordings (Baylor et ai., 1980) probably all of the
temporal integration is in the outer segment of the
photoreceptor.
The FH model as represented in equations (25)
and (26) is a linear model. That is, the sum of two
inputs L(t) + 12(0 yields a response y.~(t) + y.2(t)
which is the sum of the responses to the two inputs
presented alone - - the principle of Superposition.
However, Fuortes and Hodgkin (1964) described a
modification of the model which would help explain
observed departures from linearity, in particular the
change of gain with changes in mean level of
illumination which is the basis of photoreceptor
light adaptation. They suggested that the rate
constant of decay, a2, was increased by increase of
steady level of the output stage y, but that the rate
constant for buildup al was independent of
illumination. Thus, the scheme they proposed was:
a2 = ao2(1 +f0'.)) (27)
where the function fLy.) is chosen to fit the
dependence of gain on mean level.
The point of the FH model for adaptation is that
the gain and time course of response are thereby
linked, which may be seen as follows. Say T2 =
1/a2. The impulse response (that is, the response
to a brief flash in the linear range) of the simple
n-stage F H model is, [cf. equation (7) of Fuortes
and Hodgkin, 1964]
Q . A . T2"-'. (tlT2)"-'e -'IT2 (28)
where Q is the amount of light in the flash and A
is a constant proportional to (al)". The time to peak
o f this impulse response is ( n - 1)T2 and the gain
is proportional to /'2"-1. If only T2 is reduced by
adaptation, as assumed by Fuortes and Hodgkin,
the time to peak will shorten and the gain will drop,
much more steeply than the time to peak.