The active form of phytochrome: a new hypothesis based on phytochrome pelletability studies

Difficulties arising from the current dogma that the far-red absorbing form of phytochrome (P_fr) is the only active form are discussed.A new hypothesis is proposed in which phytochrome is held to be the photoreceptor for both low energy (pulse) and high energy (HIR) responses. There is a common basic mechanism of action involving interaction between phytochrome and a binding site within the cell. The phytochrome involvement in low energy responses exhibits an action spectrum for binding that matches the P_r absorption spectrum and reversibility by far-red irradiation. Upon prolonged irradiation the phytochrome-binding site interaction acquires different characteristics that are reminiscent of those displayed in HIR, e.g. dependence on sustained irradiation for continual binding, dependence of the degree of binding on irradiance and the similarity of the action spectrum with that of HIR action spectra, e.g. that for inhibition of lettuce hypocotyl lengthening.As expected on the basis of the new hypothesis the particulate fraction of phytochrome contains both P_r and P_fr. Arguments are advanced that the presence of P_r in pellets of particulate phytochrome cannot be accounted for by (i) the “induced fit” hypothesis, (ii) the “pigment cycling” hypothesis, and (iii) the “open phytochrome-receptor model”. We conclude that phytochrome molecules, after being sufficiently energized can interact with their intracellular binding sites irrespective of their chromophoric configuration.