Phototransduction in the fly compound eye exhibits temporal resonances and a pure time delay

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Abstract
Photoreceptor cells in both vertebrates and invertebrates respond to a flash of light with a slow graded change in membrane potential, which is generally depolarising in invertebrates and hyperpolarising in vertebrates. Although some of the early photochemical and biochemical stages of the transduction process have been elucidated in both cases, these reactions are fast compared with the time course of the electrical response, which is typically hundreds of milliseconds long. To explain this slow response in the eye of Limulus, Fuortes and Hodgkin proposed a mechanism in which the light signal passes through a cascade of simple low pass filters. The model was later defined more specifically in terms of a chain of chemical reactions linked together through products and reactants, and has been used with small modifications and different numbers of stages to account for the behaviour of various vertebrate and invertebrate photoreceptors including the fly compound eye. I have now obtained evidence that phototransduction in the fly in small signal conditions involves underdamped resonance behaviour and a significant pure time delay, neither of which can be accounted for by the conventional cascade model.