植物非编码RNA调控春化作用的表观遗传-张绍峰

HEREDITAS(Beijing)20127,34(7):829―834ISSN0253-9772综述收稿日期:2011−12−13;修回日期:2012−02−02基金项目:(30730053)(,09JC1416000)作者简介:,,E-mail:[e-mail]通讯作者:,,,E-mail:[e-mail]网络出版时间:2012-5-2202:42:27URL::10.3724/SP.J.1005.2012.00829植物非编码RNA调控春化作用的表观遗传张绍峰,李晓荣,孙传宝,何玉科,,200032在自然界中许多高等植物需要通过冬季的低温阶段实现从营养生长到生殖生长的时期转化,这一生物学过程称作春化作用。小麦(TriticumaestivumL.)和油菜(BrassicanapusL.)等作物以种子为产品器官,生产上往往通过茬口安排和栽培措施使植株尽早通过春化作用,以促进花芽形成和花器官发育,而大白菜(Brapassp.pekinenesis)和甘蓝(B.oleracea)等作物以叶球等营养器官作为产品器官,生产上则设法避免低温引起的春化作用,以保证产品器官的充分生长。FLOWERINGLOCUSC(FLC)作为一种重要的开花抑制蛋白负调控春化作用,参与植株从营养生长向生殖生长的转化过程。文章综述了春化中FLC表达受抑制主要通过低温诱导表达FLC基因区域的非编码RNA以及VRN1、VRN2、VIN3等蛋白参与介导组蛋白甲基化,从而在表观遗传上控制春化作用的进程和产品器官的正常发育。FLC;表观遗传;春化作用;非编码RNA;天然反向转录本Epigeneticsofplantvernalizationregulatedbynon-codingRNAsZHANGShao-Feng,LIXiao-Rong,SUNChuan-Bao,HEYu-KeStateKeyLaboratoryofPlantMolecularGenetics,ShanghaiInstituteofPlantPhysiology&Ecology,ShanghaiInstitutesforBiologicalSciences,ChineseAcademyofSciences,Shanghai200032,ChinaAbstract:Manyhigherplantsmustexperienceaperiodofwintercoldtoaccomplishthetransitionfromvegetativetoreproductivegrowth.Thisbiologicalprocessiscalledvernalization.Somecropssuchaswheat(TriticumaestivumL.)andoilseedrape(BrassicanapusL.)produceseedsasedibleorgans,andthereforespecialmeasuresofrotationandcultivationarenecessaryforplantstogothroughanearlyvernalizationforflowerdifferentiationanddevelopment,whereastheothercropssuchasChinesecabbage(Brapassp.pekinenesis)andcabbage(BrassicanapusL.)produceleafyheadsasedibleorgans,andadditionalpracticeshouldbetakentoavoidvernalizationforaprolongedandfullyvegetativegrowth.Beforevernalization,floweringisrepressedbytheactionofagenecalledFloweringLocusC(FLC).Thispaperreviewedthefunc-tionofnon-codingRNAsandsomeproteinsincludingVRN1,VRN2,andVIN3inepigeneticregulationofFLCduringvernalization.Keywords:FLC;epigenetics;vernalization;non-codingRNA;naturalantisensetranscripts830HEREDITAS(Beijing)201234,(TriticumaestivumL.)(BrassicanapusL.),,(Brassicarapassp.pekinenesis)(Brassicaoleracea),,,,,,,[1,2]1FLC,,FLOWERINGLOCUSC(FLC),FLCMADS,76,FT(FLOWERINGLOCUST)SOC1(SUPPRESSOROFOVEREXPRESSIONOFCO1)[3~8]FLCFLC,,(Arabidopsisthaliana)ColumbiaFLC,,FLC,FLC4[4,6~8],3[7,8]FLC,FLC,(H3K4me3),(H3K9me3,H3K27me3)[9],FLC,FLC,[3~8]VIN3(VERNALIZATIONINSENSITIVE3),FLC,,[8]H3K4me3FLD(FLOWERINGLOCUSD)[5]VRN2(VERNALISATION2)LHP1(LIKEHETEROCH-ROMATINPROTEIN1)H3K27me3H3K9me3FLC[10]RNAFLC,FLCRNARNA[11~13](H3)FLCH3K4me3(Trithorax-groupH3K4me3),;H3K27me3(Poly-comb-groupH3K27me3),,TrithoraxgroupPcG,H3K4me3H3K27me3FLCTrithorax-groupPolycomb-group,H3[14,15],,FLC,FLCH3ly4ly36ly79ly9ly14,FLC,FLCH3ly4ly9ly14,H3ly9ly27,FLC2RNAFLC,,FLCRNACOOLAIR(ColdinducedlongantisenseintragenicRNA)COLDAIR(ColdassistedintronicnoncodingRNA),FLC[11,12]7:RNA831COOLAIRFLC,mRNARNANATs(Naturalantisensetranscripts),NATsRNA,[16~18],RNA,NATsRNADNA(Genomeimprinting)[19,20],RNADCL1(DICER-LIKE1)HYL1(HYPONASTICLEAVES1)RNAnat-siRNAnat-siRNANATs,NATs,[21]NATsnat-siRNARNA,RNA,NATs,,NATs,,NATs,COOLAIR5′3′POLYARNA,POLII,FLC3′COOLAIR[11],COOLAIR(1),COOLAIR,,FLCLiu[22],23′CstF77CstF64,FLC3′,CstF(Cleavagesti-mulatoryFactor)77CstF64RNACOOLAIRPOLYA,,FLC,图1FLC基因结构及其天然反义转录本COOLAIR1FLC,,2FLCCOOLAIR,,34FLCCOOLAIR[11]RNACOLDAIR5′POLYARNA(2)FLC[12,13]COLDAIRRNAiCOLDAIR,COLDAIRPcGFLC,,,,,,FLC,,,,DNA(),(H3)[23]图2从FLC基因第一个内含子中转录生产的非编码RNACOLDAIR12COLDAIR[12]2007,FLC3’siRNA,24nt30ntsiRNA,siRNAFLC[24,25],832HEREDITAS(Beijing)201234siRNAFLC3,FLC,VIN3VRN2VRN1Trithorax-groupPolycomb-groupH3vrn1FLCH3ly9,,VRN1vrn1vrn2FLC,,vrn1vrn2FLC,,VRN1VRN2HP1LHP1,FLCH3ly9FLC[3,5,10],FLC,,FLC,VRN1VRN2VIN3LHP14,vrn1vrn2vin3lhp1,FLC,H3ly27VIN3,vin3FLCvrn1vrn2lhp1,FLCvrn2FLC;vrn1,H3ly9,H3ly27,H3ly27FLC;lhp1H3ly27ly9,FLC,FLCVIN3FLCH3ly27ly9FLC,H3ly27ly9LHP1FLCLHP1FLCFLC[3,5,10]4FLCMAF1(MADSAFFECTINGFLOWERING1)MAF2MAF3MAF4MAF5AGL19AGL24(AGAMOUS-LIKE24)7MADS,MAF1MAF2MAF3MAF4,,MAF1,MAF2MAF3MAF4MAF5MAF2MAF3MAF4MAF5,MAF2,,,MAF2AGL19AGL24FLCFLC,MAF2MAF2,SOC1FTAGL19AGL24,AGL19AGL24,MAF2AGL19FLCVIN3AGL19H3ly27H3ly9,H3ly27[26~33]5,FLC,FLC,FLC,FLC,FLC7:RNA833,FLC20073FLC,BrpFLC1BrpFLC2BrpFLC3BrpFLC22,BrpFLC110,BrpFLC33[8],20024FLC,BrpFLC1BrpFLC2BrpFLC3BrpFLC5[4,34],,,,FLC,,FLC,nat-siRNA,FLCsiRNA,DNA;FLC,,,,;FLC,,,,FLC,,参考文献(References):[1]SwiezewskiS,CrevillenP,LiuF,EckerJR,JerzmanowskiA,DeanC.SmallRNA-mediatedchromatinsilencingdirectedtothe3′regionoftheArabidopsisgeneencodingthedevelopmentalregulator,FLC.ProcNatlAcadSciUSA,2007,104(9):3633–3638.[2]GroszmannM,GreavesIK,AlbertN,FujimotoR,Helli-wellCA,DennisES,PeacockWJ.Epigeneticsinplants-vernalizationandhybrid.BiochimBiophysActa,2011,1809(8):427–437.[3]KoleC,QuijadaP,MichaelsSD,AmasinoRM,OsbornTC.Evidenceforhomologyofflowering-timegenesVFR2fromBrassicarapaandFLCfromArabidopsisthaliana.TheorApplGenet,2001,102(2–3):425–430.[4]SchranzME,QuijadaP,SungS,B,LukensL,AmasinoR,OsbornTC.CharacterizationandeffectsofthereplicatedfloweringtimegeneFLCinBrassicarapa.Genetics,2002,162(3):1457–1468.[5]KimSY,MichaelsSD.SUPPRESSOROFFRI4