Far-red light blocks greening of Arabidopsis seedlings via a phytochrome A-mediated change in plastid development

We have characterized a far-red-light response that induces a novel pathway for plastid development in Arabidopsis seedlings. This response results in the inability of cotyledons to green upon subsequent white light illumination, and the response is suppressed by exogenous sucrose. Studies with mutants showed that this far-red block of greening is phytochrome A dependent and requires an intact downstream signaling pathway in which FHY1 and FHY3 may be components but in which HY5 is not. This highlights a previously undefined branchpoint in the phytochrome signaling pathway. Ultrastructural analysis showed that the far-red block correlates with both the failure of plastids to accumulate prolamellar bodies and the formation of vesicles in the stroma. We present evidence that the far-red block of greening is the result of severe repression of protochlorophyllide reductase (POR) genes by far-red light coupled with irreversible plastid damage. This results in the temporal separation of phytochrome-mediated POR; repression from light-dependent protochlorophyllide reduction, two processes that normally occur in coordination in white light.