Longitudinal spread of adaptation in the rods of the frog's retina.

1. The stimulus-response function of the red rods in the retina of the common frog (Rana temporaria) was determined in different adaptational states by measuring aspartate-isolated receptor responses. 2. Flash stimuli, background adaptations and bleaches were delivered through the same optical channel forming an oblique light-beam striking the receptor side of the isolated and flat-mounted retina at an angle of 10 degrees. 3. When the light was blue-green and optimally polarized the absorbance of the receptor layer was about 2, from which follows that 70-80% of the light was absorbed in the distal third of the rod outer segments, i.e. the exposure was local. Homogeneous exposures of the whole rod outer segments were obtained with orange and red lights. 4. Combinations of homogeneous and local stimuli with homogeneous and local adaptations were used to investigate the longitudinal spread of background, intermediate and opsin adaptation, i.e. the sensitivity-reducing effect of a background light, and the transient and permanent sensitivity losses following a bleach isomerizing 3.5-26% (usually 10%) of the rhodopsin in the retina. 5. The results obtained were related to predictions based both on the assumption that the adaptation effects spread longitudinally within the rod outer segments and the assumption that they are strictly confined to the disks absorbing the adapting lights. 6. These comparisons reveal that all three types of adaptation spread longitudinally. It is for instance clear that the sensitivity loss observed with homogeneous stimuli and local adaptation (as compared to homogeneous adaptation) is larger than that predicted by the non-spreading hypothesis. 7. The longitudinal spread of background adaptation is largely finished within 10 sec after turning on the background light, while an efficient spread of the intermediate adaptation effect may require minutes. 8. A background light decreasing the sensitivity by about one log unit decreases the time from flash to response maximum from 5 to 1 sec (small responses). Corresponding opsin adaptation effects are accompanied by less dramatic changes in response kinetics. 9. Independent of adaptation type - homogeneous or local, background, intermediate or opsin - it was found that local stimuli are less efficient that homogeneous stimuli in light-adapted retinae. This effect can be explained assuming that the sensitivity-reducing effects are pronounced in the distal than in the proximal parts of the rod outer segments. 10. The opsin adaptation effect following 10% local bleaches decreases the sensitivity to both homogeneous and local stimuli 2-3 times more than corresponding homogeneous bleaches. This means that the strength of the opsin effect is not related to the average percentage bleached but to the fraction bleached in the distal part of the rod, or generally to the fraction bleached in the most affected region. 11...