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Duplicity Theory of Vision: From Newton to the Present

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158 <strong>the</strong>ories <strong>of</strong> sensitivity regulationlight did not raise <strong>the</strong> quantum demand at <strong>the</strong> recep<strong>to</strong>r level (i.e.<strong>the</strong> threshold for rod-recep<strong>to</strong>r signals did not increase) and that alladaptation occurred at <strong>the</strong> pool.This conclusion was also supported by experiments on darkadaptation where <strong>the</strong> bleaching light was sent by means <strong>of</strong> a briefelectronic flash ei<strong>the</strong>r through a grating <strong>of</strong> 0.5º period <strong>to</strong> give astriped bleach or through a neutral filter <strong>to</strong> pass an equal amount <strong>of</strong>light spread evenly on <strong>the</strong> retina. The test flash was under <strong>the</strong> lattercondition a grating <strong>of</strong> 0.5º period, while it was an equalizing neutraltest light in <strong>the</strong> former. Again, no significant differences between <strong>the</strong>two conditions were obtained.Rush<strong>to</strong>n (1965a) argued that if sensitivity regulation occurredat <strong>the</strong> recep<strong>to</strong>r level, <strong>the</strong> dark bars <strong>of</strong> <strong>the</strong> grating should have shielded<strong>the</strong> retina from <strong>the</strong> flash <strong>of</strong> <strong>the</strong> bleaching light. The retina should thushave been fully dark adapted at <strong>the</strong> shielded regions, while every rodin <strong>the</strong> bleached area should have been substantially bleached when<strong>the</strong> same <strong>to</strong>tal bleaching light was evenly distributed on <strong>the</strong> retina.Hence, from his results he concluded that bleaching with <strong>the</strong> gratingor with <strong>the</strong> same light spread evenly had about <strong>the</strong> same effect on<strong>the</strong> threshold level, and also that <strong>the</strong> threshold level obtained by <strong>the</strong>test flash was <strong>the</strong> same whe<strong>the</strong>r <strong>the</strong> test energy was concentratedby <strong>the</strong> grating or spread evenly – all in harmony with his AGC-poolhypo<strong>the</strong>sis.20.2 Ana<strong>to</strong>mical and electrophysiologicalevidenceAna<strong>to</strong>mical and electrophysiological evidence supporting Rush<strong>to</strong>n’spost-recep<strong>to</strong>r adaptation <strong>the</strong>ory was presented by Dowling (1967).Toge<strong>the</strong>r with Boycott he had examined <strong>the</strong> synaptic organization<strong>of</strong> <strong>the</strong> retina in several types <strong>of</strong> vertebrate retina, including <strong>the</strong>primate, using an electron microscope. Striking support for Rush<strong>to</strong>n’sAGC-pool hypo<strong>the</strong>sis was obtained. Thus, <strong>the</strong>y found widespreadreciprocal synapses between bipolar cell terminals and amacrine-cellprocesses. The ana<strong>to</strong>mical evidence indicated that bipolar cells made

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