Rhodopsin and visual adaptation: analysis of photoreceptor thresholds in the isolated skate retina.

Photoreceptor thresholds in the isolated retina of the skate, determined by extracellular measurement of the photoreceptor potential during periods of light and dark adaptation, were analyzed in relationship to prevailing states of the visual pigment. The starting assumption of the analysis is that relative levels of three forms of the pigment molecule [native rhodopsin (R), a photoactivated intermediate (R*), and bleached pigment (B)] govern (quasi-) stable levels of threshold measured (a) during exposure of the retina to background light of fixed incident intensity (Ib), and (b) after irradiation that bleaches a defined fraction (B) of the rhodopsin. It is shown that experimental data are described well by the equation It/ It0 = (1 - B)-1 X F X (1 + 0(3)B), where F = [1 + 0(1)Ib(1 - B) + 0(2)B]. In this equation, It/ It0 is the relative threshold for detection of a test flash; (1 - B) approximates the relative efficiency of quantum capture; and 0(1) - 0(3) are constants. For values of 0(1) - 0(3) yielding an optimal fit to experimental data, log (It/ It0 ) approximately log F over a broad range of values of Ib and B. It is further shown that the algebraic form of the term F in the above equation is consistent with the predictions of a (steady-state) model for the role of the pigment molecule in photoreceptor adaptation. The model proposes that R* and B desensitize the photoreceptor by acting (in qualitatively similar fashion) to reduce the availability of E, an intracellular substance whose activation supports generation of the flash response. Results of the analysis are discussed in relation to the Dowling- Rushton equation (Dowling, 1960, 1963; Rushton , 1961), and to the results of more recent studies examining light and dark adaptation.