RNA-Silencing Enzymes Pol IV and Pol V in Maize More than one Flavor 英文参考文献.docVIP

Perspective RNA-SilencingEnzymesPolIVandPolVinMaize:More thanoneFlavor? CraigS.Pikaard1,2,3*,SarahTucker3 1DepartmentofBiology,IndianaUniversity,Bloomington,Indiana,UnitedStatesofAmerica,2DepartmentofMolecularandCellularBiochemistry,IndianaUniversity, Bloomington,Indiana,UnitedStatesofAmerica,3DepartmentofBiology,WashingtonUniversityinSt.Louis,St.Louis,Missouri,UnitedStatesofAmerica There are some puzzling genetic phe- nomena in nature that have kept geneti- cistsbusilysearchingforcluesfordecades. Amonggenetics’X-filesarecasesinwhich alleles of the same gene have the same sequence but very different functional states. Gene behaviors of this sort fall undertheheadingof‘‘epigeneticphenom- ena.’’ Traits attributable to epigenetically altered alleles (epialleles) are often not transmitted between generations as inde- pendent,immutablecharacters,according totherulesthatMendelworkedoutwith his garden peas. Instead, there are cases whereallelescanchangestateifandwhen they find themselves in the same nucleus asepiallelesthatarelockedintoadifferent functional state. This is the case in paramutation, which describes the situa- tionwherebyasilent,paramutagenicallele can influence the expression of an active, paramutablealleleofthesamegenewhen thetwoallelesareexposedtooneanother within the same genome [1]. The silent allele, which is defined as being paramu- tagenic, somehow transfers its silent state totheotherallele,causingthatpreviously activealleletobecomesilenced,too.Most interestingly,thepreviouslyactivealleleis nowstablyalteredinsuchawaythatitcan retain the silenced state through meiosis and future generations. Moreover, the newly silenced allele is paramutagenic itselfandwillsilencecompatible paramu- table alleles. All of this happens without anychangesinthenucleotidesequenceat the affected locus. This unusual allelic behavior is the genic analogue of the shape-shiftingthatoccursamongproteins thatcanconverttoaprionstateandthen cause other proteins of identical