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romes in the cell as that would provide clues as to their mode of action in the 1980s lee pratt s lab at the university of georgia had used antibodies directed principally against phya to label it within the cell finding that whereas pr was distributed throughout the cytoplasm photoconversion to pfr lead to rapid sequestration of the protein to small areas followed by its disappearance 77 what these sequestered areas represent has not been established the picture changed dramatically in 1999 however when workers in japan and germany using transgenic phytochromes tagged with fluorescent moieties showed that both phya and phyb as pfr are imported into the nucleus where they form speckles currently thought to be associated with signalling and or proteolysis 78 79 nuclear localisation of pfr fitted well with the reasonable idea that phytochrome acts by regulating gene expression indeed at about the same time weimin ni in the quail lab in berkeley identified the pif family of p hytochrome i nteracting transcription f actors that are thought to be the primary route of phytochrome signalling see 14 as pif3 was initially identified through its binding to a c terminal phyb fragment comprising the pas repeat and histidine kinase like modules 80 it was a great surprise to find that the primary interaction is instead with the pfr state of photosensory module 81 indeed tomonao matsushita in the nagatani lab in kyoto showed that the phyb photosensory module alone when dimerised in the nucleus functions at least as effectively as the complete phyb molecule 82 despite some initial confusion it was established that the pifs in particular pif3 act as regulators of genes responsible for the etiolated developmental form skotomorphogenesis of seedlings grown in darkness partly through production of phytohormones such as auxin that enhance stem extension this was made clear by the creation of pifq a multiple pif gene knockout mutant pifq seedlings grown in total darkness developed similarly to wild type seedlings grown in the light generally speaking pifs promote skotomorphogenesis and repress photomorphogenesis solving the 3d crystal structures of the prokaryotic phytochrome photosensory module see above encouraged similar studies of plant phytochromes the first success was in 2014 for the photosensory module of arabidopsis phyb pr by sethe burgie in the vierstra lab 83 pdb 4our equivalent pr structures of other plant phytochromes at higher resolution were published subsequently 26 although the sequence identity relative to the photosensor of cph1 is only about 30 the 3d structures are remarkably similar as in the case of prokaryotic phytochromes it has not been possible to crystallise a complete plant phytochrome but the emergence of cryo electron microscopy cryo em allowed near complete 3d structures of phya pdb 8f5z and others and phyb pdb 7rzw in the pr state to be solved by single molecule imaging how light and thus pfr formation regulates 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