Recovery kinetics of the saturating photocurrent
response in amphibian rods suggest regulation of the visual
signal by a first-order deactivation reaction with an exponential
time constant (τc) of about 2 s. The original
hypothesis that τc represents the lifetime
of activated rhodopsin (R*) in a single-step
deactivation appears at odds with several recent findings,
for example, that Ca2+, a known regulator of
the enzymatic phosphorylation of R*, does not
regulate the value of τc. A recently proposed
alternative hypothesis, that τc is the lifetime
of activated transducin and that the R* lifetime
is relatively short (∼0.4 s), appears consistent with
the Ca2+ data but is difficult to reconcile
with a high specific catalytic activity of R*.
The present theoretical study proposes a rate-equation
model of R* activation and deactivation in amphibian
rods that is generally consistent with observed properties
of the τc-associated reaction and the action
of Ca2+ as well as with the stereotyped nature
of the single-photon response. The model is developed by
considering the effect of background light on a time-dependent
variable, Reff*,
defined as the effective total level of R*
activity. Central starting assumptions are that Ca2+
reduction mediates the effect of background light on Reff*(t) and
that background desensitization of the photocurrent flash
response derives from this action of Ca2+. Construction
of the model is guided by criteria based on previous experimental
findings. Among these are the approximate constancy of
background desensitization expressed at near-peak and later
times in the flash response, and the large (∼10-fold)
dynamic range of this desensitization. The proposed model
hypothesizes that an event regulated by Ca2+
feedback causes activated rhodopsin to become susceptible
to a two-phase, stochastic deactivation process, the second
phase of which is characterized by τc. A
central prediction of the model is the regulated transition
of flash-activated R* to “R**”,
a state exhibiting greatly increased catalytic activity.