文獻(xiàn)綜述樹木次生木質(zhì)部的形成及其分子調(diào)控機(jī)制

日菊進(jìn)行細(xì)胞培養(yǎng)，對影響導(dǎo)管組織分化、次生細(xì)胞壁的合成及細(xì)胞程序性過程的調(diào)節(jié)因子進(jìn)行了鑒定；鑒于擬南芥具有有利于生長發(fā)育研究的獨(dú)特的資源——根部在擬南芥中利用操控技術(shù)及轉(zhuǎn)錄組學(xué)，對調(diào)控木材形成的關(guān)鍵因子進(jìn)行了鑒定(Schraderetal.,2004)，同時驗(yàn)證了楊樹中調(diào)控形成層活性及部分木質(zhì)化的轉(zhuǎn)錄因子和激素的功能，并對激素與間的相互作用進(jìn)行了探究。研究結(jié)果表明，楊樹和擬南芥中Shrdr(2004)和Gora(2011)等人利用較空間分辨率的轉(zhuǎn)錄組學(xué)和化學(xué)分析方法對楊樹樹干的木材形成區(qū)域進(jìn)行了研究此外現(xiàn)代技術(shù)的發(fā)展以及樹木組豐富的遺傳多樣性使得樹木遺傳學(xué)得到快速發(fā)展到目前為止已得到一些森林樹種及針葉樹的組如楊樹桉樹(Grttpglia,tal.,2012)白云杉(Nytedt,tl,2013)。云杉組的發(fā)布為研究木材形成過程提供了有力的平臺方便了研究者們對被子植物及子植物全組進(jìn)行比較研究被子植物和子植物的一大差別在于兩者的木質(zhì)部：子植物的木質(zhì)部僅由管胞組成而被子植物既有導(dǎo)管組織(導(dǎo)管和管胞)又包含了纖維物質(zhì)。這也表明了在木質(zhì)部中可能存在這兩種不同的細(xì)胞類型。D(VSUAREATEDNC gnes)被證實(shí)在木質(zhì)部分化中起到主要的調(diào)節(jié)作用。在云杉組中，只發(fā)現(xiàn)了兩種D，而在擬南芥中則發(fā)現(xiàn)了7種。這一結(jié)果表明D的擴(kuò)增在被子植物木質(zhì)部的形成過程中起到重要作用(Nytedt,etal.,2013)以及棕櫚樹等(Peng,etal2013Singh,etal2013,Al-Mssallem,etal2013)不斷增大。這些資源為我們研究的演變規(guī)律(這對于研究形成層的起始反應(yīng)和作為組學(xué)研究的補(bǔ)充，樹木遺傳學(xué)的研究近期取得了突破性進(jìn)展：使樹木結(jié)構(gòu)發(fā)生特異性突變的首個得到了鑒定。通過對后代繪制圖譜，Dardick等人(2013)發(fā)現(xiàn)PpeTAC1(TILLERANGLECONTROL1)可以調(diào)控桃樹樹干的分枝角度。該的半顯性突變使得桃樹樹干有了水平、垂直和柱形的性狀分離。圖譜的成功繪制再次肯定了發(fā)現(xiàn)調(diào)控樹木各種性狀的的可能性，這為被子植物木質(zhì)化的調(diào)控機(jī)有研究，TDIF/CLE41/CLE44-TDR/PXY-WOX4對形成層活性的維持進(jìn)行調(diào)控(Hirakawa,etal.,2010,Etcs,etal.,2010)。在擬南芥中，CLE41/CLE44的翻譯產(chǎn)物在韌皮部經(jīng)加工后得到導(dǎo)管分化抑制因子TDIF，TDIF與其受體TDR/PXY相結(jié)合。WOX4(WUSHCHEL–likeHOMEOBOX4)，可以通過調(diào)節(jié)TDIF與受體的結(jié)合來調(diào)控形成層細(xì)胞或分化。wox4突變體的下胚軸和花莖部位，形成層的活性降低，這表明WOX4在調(diào)控分生組織活動中起到一定的作用(Hirakawa,etal.,2010,Suer,etal.,2011)。WOX14同樣能夠起到調(diào)控形成層細(xì)胞增殖的作用。研究表明，TDIF/CLE41/CLE44-TDR/PXY-WOX4信號通路無論在木本植物還是草本植物進(jìn)化過程中似乎都是保守的，另有研究，ERECTA(ER)/ER-LIKE1及EPFL4/6可能會對TDR/PXY產(chǎn)生影響(Etcs,etal.,2013,UchidaandTasaka,2013),它們的變異可能會增加在維管結(jié)構(gòu)中tdr/pxy突變體的比例。另外，MOL1(MORELA LGROWTH1)和RUL1(REDUCEDINLA LGROWTH1)極有可能在TDR/PXY-WOX4通游表達(dá)，這兩種受體蛋白激酶在形成層活性調(diào)控中起到相反的作用，MOL1對花莖的次生生長起到抑制作用，而RUL1可促進(jìn)花莖的次生生長(Agusti,etal.,2011)。KNOXKNOX質(zhì)部的分化則受到抑制。在生物中，BP突變型比野生型的木質(zhì)部和纖維細(xì)胞木質(zhì)化時間更早。這表明了KNOX在木質(zhì)部次生分化過程中起到一定的作用(Du,MansfieldandGroover2004)BP突變體中同樣發(fā)現(xiàn)了異常的木質(zhì)素沉積現(xiàn)象(Mele,etal.,2003)。要激一。在楊樹中，通過過表達(dá)突變的IAA3，可以阻斷生長素信號通路，抑制形成層細(xì)胞的擴(kuò)增，從而減少木材的形成(Nilsson,etal.,2008)。TDR/PXY-WOX4在生長素信號通路的下游表達(dá)，可以調(diào)控形成層細(xì)胞的增殖(Suer,etal.,2011)。細(xì)胞分在缺乏細(xì)胞素生物合成酶IPT的擬南芥突變體ipt1,3,5,7中，次生生長受到了抑制(Matsumoto-Kitano,etal.,2008)。反之，細(xì)胞素的濃度提高，次生生長會得到加強(qiáng)。形成層的細(xì)胞增殖(Loveetal2009)；在缺乏TDR/PXY通路的擬南芥之中，ERF109和ERF018(ETHYLENEINDUCEDRESPONSEFACTORgenes)雙突變的的次生生長受到抑制，而乙烯過表達(dá)的突變體的生長則有所增強(qiáng)(Etcs,ProvostandTurner,2012)。近年來，楊樹ERF全圖譜得到建立，證實(shí)了ERF具有調(diào)控次生生bHLH(basichelix-loop-helix)TF二聚體是調(diào)控維管組織早期發(fā)育的重要因子，主要包括TMO5(TARGETOFMONOPTEROS5)，LHW(LONESOMEHIGHWAY)以及它們的同源物(DeRybeletal.,2013)。在TMO四倍TMO5及LHW的的表達(dá)后，根部維管組織中重新實(shí)現(xiàn)了大量的平周。有研究，在擬南芥根部維管組織發(fā)育過程中，存在乙烯與生長素共同調(diào)控成層活性和木質(zhì)部形成的機(jī)制(Bishopp,etal.,2011)，編碼亮氨酸拉鏈(HD-ZIPIII)在初生生長過程中對木質(zhì)部規(guī)格起到重要調(diào)控作化(Carlsbeckeretal2010)。HD-ZIPIII的水平受到SHR及miR165/166的調(diào)節(jié)，從而調(diào)控根部原生木質(zhì)部的分化(Carlsbecker,etal.,2010)。油菜素內(nèi)酯(BR)ZIPIII的表達(dá)，從而促進(jìn)木質(zhì)部的分化(Ohashi-ItoandFukuda,2003)。楊樹中的POPCORONAmiRNA有抑制作用，POPCORONA的沉默可導(dǎo)致髓細(xì)胞的異常木質(zhì)POPCORONA的過表達(dá)則可使木質(zhì)部及韌皮纖維的木質(zhì)化延遲(Du,etal2011)。miRNA抑制物過表達(dá)時，會使得皮層薄壁組織內(nèi)極性顛倒，形成層層次異位(Robischon,etal.,2011)。NAC轉(zhuǎn)錄因子被證實(shí)是木質(zhì)部分化的主要調(diào)控因子。VND7誘導(dǎo)原生木質(zhì)部分化，VND6可誘導(dǎo)后生木質(zhì)部分化。SND1/NST3以及NST1都可以促進(jìn)纖維分化。在楊樹中，PtVNS/PtrWND的過表達(dá)可誘導(dǎo)異常的木質(zhì)分化(Ohtani,etal.,2011)，而VNI2VND7的表達(dá)(Yamaguchi,etal.,2010)。楊樹中的半乳葡甘露聚低聚糖(GGMOs)已被證實(shí)可抑制NAC的表達(dá)(Zhao,etal.,2013)，因此，內(nèi)源甘露聚糖酶PtrMAN6的過表達(dá)可抑制次生細(xì)胞壁的增厚，沉默時則無法起到抑制作用(Zhao,etal.,2013)。TDIF可促進(jìn)樹干形成層細(xì)胞增殖，卻對葉片及下胚軸的維管分化有著抑制作用可通過激化細(xì)胞素相關(guān)通路來抑制根部木質(zhì)部維管組織分化(Kondo,etal.,2011)。形成(MauriatandMoritz,2009)。NAC可通過誘導(dǎo)兩個MYB轉(zhuǎn)錄因子——MYB46和MYB83的表達(dá)來啟動木質(zhì)化程序(Zhong,RichardsonandYe2007McCarthy,ZhongandYe2009)。起到這一作用VND6、VND7、NST1SND1/NST3。MYB46/83節(jié)點(diǎn)激發(fā)了其他轉(zhuǎn)錄因子RichardsonandYe,2007,McCarthy,ZhongandYe,2009,ZhongandYe,2012)，它們在促總的來說，木質(zhì)部分化是一個極其復(fù)雜的過程，NAC及MYB，可誘導(dǎo)許多相關(guān)基及MYB的直系同源物現(xiàn)也已在楊樹、桉樹及松樹等樹種中得到了發(fā)現(xiàn)和鑒定LeeandYe,2010,Zhong,etal.,2013)木質(zhì)部各類細(xì)胞的次生細(xì)胞壁上的沉積物主要取決于周質(zhì)微管(corticalmicrotubules，MT)的作用。周質(zhì)微管直接決定了纖維素合酶復(fù)合物的動態(tài)(Paredez,SomervilleandEhrhardt,2006)MT關(guān)聯(lián)蛋白(MT-ASSOCIATEDPROTEIN70，MAP70)的過表達(dá)可提高細(xì)胞壁螺紋的出現(xiàn)頻率，而它們的沉默表達(dá)則會導(dǎo)致有凹痕的細(xì)胞壁的出現(xiàn)(Pesquet,2010)。相比之下，MT解聚蛋白MIDD1MT的解聚合水平，從而在細(xì)胞壁的一些區(qū)域形成凹痕；在降解MIDD1后，又重新出現(xiàn)了了無凹痕的細(xì)胞壁(Oda,2010)。MIDD1在質(zhì)膜的固定是通過ROPGTP酶調(diào)控而實(shí)現(xiàn)的，在質(zhì)膜上，ROP11可刺激MIDD1在形成凹痕的部位起作用(OdaandFukuda,2012)。細(xì)胞程序性(ProgrammedCelldeath，PCD)是木質(zhì)分化的最后一個階段。在高等植物中，導(dǎo)管分子分化及木質(zhì)部形成過程包括一系列的細(xì)胞結(jié)構(gòu)和物質(zhì)的合成。PCD過程中，液泡的對木質(zhì)部分化至關(guān)重要。當(dāng)液泡破裂后，其中核酸酶和蛋白酶纖維的降解則緩慢一些，在液泡破碎之前，纖維細(xì)胞內(nèi)DNA發(fā)生，高度凝集，細(xì)胞逐漸分解(Courtois-Moreau,etal.,2009)。在楊樹次生木質(zhì)部形成過程中，20SP(20Sproteasome)20SP的作用，楊樹和擬南芥導(dǎo)管系統(tǒng)的PCD階段均可遭到破壞(Han,2012。XCP1(XYLEMCYSTEINEPROTEASE1)和XCP參與了導(dǎo)管系統(tǒng)的PCD過程，VND6可誘導(dǎo)這兩種蛋白酶的表達(dá)(Avci2008)。另外一種半胱氨酸蛋白酶AtMC9被證實(shí)在擬南芥的維管組織PCD(Bollhoner,2013)PCD發(fā)生的時間受到ACL5(ACUALIS5)的調(diào)控，在楊樹中的研究表明，ACL5的異常過表達(dá)會延遲楊樹的木質(zhì)部分化成時間(Milhinhos,2013)。液泡中的木質(zhì)素單體釋放出木質(zhì)素，并在導(dǎo)管細(xì)胞壁及纖維細(xì)胞中不斷沉積(Pesquet進(jìn)行ParedezSomervilleandEhrhardt2006,Pesquet,etal2013)。miRNA對木質(zhì)化的調(diào)節(jié)也有一定的作用。研究表明，Ptr-miR397a參與了楊樹漆酶的轉(zhuǎn)錄后調(diào)節(jié)過程(Luetal.,2013)。近期由科研工作者發(fā)布了竹子和棕櫚類植物的組序列，這讓我們有機(jī)會探究“木質(zhì)化的”是保守的竹子組與楊樹組一樣控制細(xì)胞壁組成及結(jié)構(gòu)的重復(fù)，如控制纖維素、木質(zhì)素生物合成酶形成的(Peng,etal.,2013)。對于TDIF/CLE41/CLE44-TDR/PXY-WOX4信號通路中“多肽-受體-靶”各組wox4突變體另外在楊樹ipt1,3,5,7突變體中未發(fā)現(xiàn)次生生長故而找到在細(xì)胞素信號通路由一個包含各種轉(zhuǎn)錄因子的調(diào)控網(wǎng)絡(luò)(GRN)調(diào)控的。在以前的研究中，曾通過酵母菌單雜交和雙雜交系統(tǒng)來在挑選出來的諸多轉(zhuǎn)錄因子中研究蛋白質(zhì)之間的相互作用生物進(jìn)程(Xu,etal.,2013)。在一個模擬的生物實(shí)驗(yàn)中，107個保守的維管組得到了鑒定，這些組很可能構(gòu)成一個復(fù)雜的具有多功能連接的聯(lián)合表達(dá)網(wǎng)絡(luò)。將全組技術(shù)與計(jì)算機(jī)模擬分析、相結(jié)合，讓我們相信，調(diào)控木質(zhì)形成的重要GRN將GRN，并將為以木材為目標(biāo)的樹木育種提供理在綜述完成過程中，老師為我們提供了大量資料和悉心指導(dǎo)，并對 AgustiJ.,LichtenbergerR.,SchwarzM.,etal.CharacterizationoftranscriptomeremodelingduringcambiumformationidentifiesMOL1andRUL1asregulatorsofsecondarygrowth[J].PlosGenetics,2011,7: Al-MssallemI.S.,HuS.,ZhangX.,etal.GenomesequenceofthedatepalmPhoenixdactyliferaL[J].NatureCommunications,2013,4:2274.AvciU.,PetzoldH.E.,IsmailI.O.,etal.CysteineproteasesXCP1andXCP2aidmicro-autolysiswithintheintactcentralvacuoleduringxylogenesisinArabidopsisroots[J].ntJournal,2008,56:303-315.BollhonerB.,ZhangB.,StaelS.,etal.PostmortemfunctionofAtMC9inxylemvesselelements[J].NewPhytologist,2013,200:498-510.BirolI.,RaymondA.,JackmanS.D.,etal.Assemblingthe20Gbwhitespruce(Piceaglauca)genomefromwhole-genomeshotgunsequencingdata[J].Bioinformatics,2013,29:1492-1497.BishoppA.,HelpH.,El-ShowkS.,etal.Amutuallyinhibitoryinctionbetweenauxinandcytokininspecifiesvascularpatterninroots[J].CurrentBiology,2011,21:917-926.BradyS.M.,OrlandoD.A.,LeeJ.Y.,etal.Ahigh-resolutionrootspatiotemporalmaprevealsdominantexpressionpatterns[J].Science,2007,318:801-806.BradyS.M.,ZhangL.,MegrawM.,etal.Ase-enrichedgeneregulatorynetworkintheArabidopsisroot[J].MolecularSystemsBiology,2011,7:459.CarlsbeckerA.,LeeJ-Y.,RobertsC.J.,etal.CellsignallingbymicroRNA165/6directsgenedosedependentrootcellfate[J].Nature,2010,465:316-321.CichanM.A.,TaylorT.N..VascularcambiumandwooddevelopmentSphenophyllum:acarboniferousarthrophyte[J].IAWABulletin,1982,3:155-Courtois-MoreauC.L.,PesquetE.,SjodinA.,etal.AuniqueprogramforcelldeathinxylemfibersofPopulusstem[J].ntJournal,2009,58:260-274.DardickC.,CallahanA.,HornR.,etal.PpeTAC1npromotesthehorizontalgrowthofbranchesinpeachtreesandisamemberofafunctionallyconservedgenefamilyfoundindiversentsspecies[J].ntJournal,2013,75:618-630.DeRybelB.,MollerB.,YoshidaS.,etal.AbHLHcomplexcontrolsembryonicvasculartissueestablishmentandindeterminategrowthinArabidopsis[J].DevelopmentalCell,2013,24:426-437.DuJ.,MansfieldS.D.,GrooverA.T..ThePopulushomeoboxgeneARBORKNOX2regulatescelldifferentiationduringsecondarygrowth[J].ntJournal,2009,60:DuJ.,MiuraE.,RobischonM.,etal.ThePopulusclassIIIHDZIPtranscriptionfactorpopcoronaaffectscelldifferentiationduringsecondarygrowthofwoodystems[J].PLoSOne,2011,6:e17458.EtcsJ.P.,ProvostC.M.,MishraL.,etal.WOX4andWOX14actdownstreamofthePXYreceptorkinasetoregulatentvascularproliferationindependentlyofanyroleinvascularorganisation[J].Development,2013,140:2224-2234.EtcsJ.P.,ProvostC.M.,TurnerS.R..ntvascularcelldivisionismaintainedbyaninctionbetweenPXYandethylenesignalling[J].PlosGenetics,2012,8: EtcsJ.P.,TurnerS.R..ThePXY-CLE41receptorligandpairdefinesamultifunctionalpathwaythatcontrolstherateandorientationofvascularcelldivision[J].Development,2010,137:767-774.GorzsasA.,StenlundH.,PerssonP.,etal.CellspecificchemotyandmultivariateimagingbycombinedFT-IRmicrospectroscopyandorthogonalprojectionstolatentstructures(OPLS)ysisrevealsthechemicallandscapeofsecondaryxylem[J].ntJournal,2011,66:903-914.GrattapagliaD.,VaillancourtR.E.,ShepherdM.,etal.ProgressinMyrtaceaegeneticsandgenomics:eucalyptusasthepivotalgenus[J].TreeGeneticsGenomes,2012,8:HanJ-J.,LinW.,OdaY.,etal.Theproteasomeisresponsibleforcaspase-3-likeactivityduringxylemdevelopment[J].ntJournal,2012,72:129-141.HejatkoJ.,RyuH.,KimG.T.,etal.Thehistidinekinasescytokinin-independent1andArabidopsishistidinekinase2and3regulatevasculartissuedevelopmentinArabidopsisshoots[J].ntCell,2009,21:2008-2021.HirakawaY.,KondoY.,FukudaH..TDIFpeptidesignalingregulatesvascularstemcellproliferationviatheWOX4homeoboxgeneinArabidopsis[J].ntCell,2010,22:2618-2629.IsraelssonM.,SundbergB.,MoritzT..Tissue-specificlocalizationofgibberellinsandexpressionofgibberellin-biosyntheticandsignalinggenesinwood-formingtissuesinaspen[J].ntJournal,2005,44:494-504.ItoY.,NakanomyoI.,MotoseH.,etal.Dodeca-CLEpeptidesassuppressorsofntstemcelldifferentiation[J].Science,2006,313:842-845.KondoY.,HirakawaY.,KieberJ.J.,etal.CLEpeptidescannegativelyregulateprotoxylemvesselformationviacytokininsignaling[J].ntandCellPhysiology,2011,52:37-48.LoveJ.,BjorklundS.,VahalaJ.,etal.EthyleneisanendogenousstimulatorofcelldivisioninthecambialmeristemofPopulus[J].ProceedingsofTheNationalAcademyofSciencesofTheUnitedStatesofAmericaUSA,2009,106:5984-LuS.,LiQ.,WeiH.,etal.Ptr-miR397aisanegativeregulatoroflaccasegenesaffectinglignincontentinPopulustrichocarpa[J].ProceedingsofTheNationalAcademyofSciencesofTheUnitedStatesofAmerica,2013,110:10848-10853.Matsumoto-KitanoM.,KusumotoT.,TarkowskiP.,etal.Cytokininsarecentralregulatorsofcambialactivity[J].ProceedingsofTheNationalAcademyofSciencesofTheUnitedStatesofAmerica,2008,105:20027-20031.MauriatM.,MoritzT..ysesofGA20ox-andGID1-overexpressingaspensuggestthatgibberellinsytwodistinctrolesinwoodformation[J].ntJournal,2009,58:989-1003.McCarthyR.L.,ZhongR.,YeZ-H..MYB83isadirecttargetofSND1andactsredundantlywithMYB46intheregulationofsecondarycellwallbiosynthesisinArabidopsis[J].ntandCellPhysiology,2009,50:1950-1964.MeleG.,OriN.,SatoY.,etal.Theknotted1-likehomeoboxgeneBREVIPEDICELLUSregulatescelldifferentiationbymodulatingmetabolicpathways[J].GenesDevelopment,2003,17:2088-2093.MilhinhosA.,PreseJ.,BollhonerB.,etal.Thermosperminelevelsarecontrolledbyanauxin-dependentfeedbackloopmechanisminPopulusxylem[J].ntJournal,2013,75:685-698.NieminenK.,ImmanenJ.,LaxellM.,etal.Cytokininsignalingregulatesdevelopmentinpor[J].ProceedingsofTheNationalAcademyofSciencesofTheUnitedStatesofAmerica,2008,105:20032-20037.NilssonJ.,KarlbergA.,AnttiH.,etal.Dissectingthemolecularbasisoftheregulationofwoodformationbyauxininhybridaspen[J].ntCell,2008,20:843-855.NystedtB.,StreetN.R.,WetterbomA.,etal.TheNorwaysprucegenomesequenceandconifergenomeevolution[J].Nature,2013,497:579-584.OdaY.,FukudaH..InitiationofcellwallpatternbyaRho-andmicrotubule-drivensymmetrybreaking[J].Science,2012,337:1333-1336.OdaY.,IidaY.,KondoY.,etal.Woodcell-wallstructurerequireslocal2D-microtubuledisassemblybyanovelsmamembrane-anchoredprotein[J].CurrentBiology,2010,20:1197-1202.Ohashi-ItoK.,OdaY.,FukudaH..Arabidopsisvascular-relatedNAC-6directlyregulatesthegenesthaternprogrammedcelldeathandsecondarywallformationduringxylemdifferentiation[J].ntCell,2010,22:3461-3473.Ohashi-ItoK.,FukudaH..HD-ZipIIIhomeoboxgenesthatincludeanovelmember,ZeHB-13(Zinnia)/ATHB-15(Arabidopsis),areinvolvedinprocambiumandxylemcelldifferentiation[J].ntandCellPhysiology,2003,44:1350-1358.OhtaniM.,NishikuboN.,XuB.,etal.ANACproteinfamilycontributingtotheregulationofwoodformationinpor[J].ntJournal,2011,67:499-512.ParedezA.R.,SomervilleC.R.,EhrhardtD.W..Visualizationofcellulosesynthasedemonstratesfunctionalassociationwithmicrotubules[J].Science,2006,312:PengZ.,LuY.,LiL.,etal.Thedraftgenomeofthefast-growingnon-timberspeciesmosobamboo(Phyllostachysheterocycla)[J].NatureGenetics,2013,45:PesquetE.,KorolevA.V.,CalderG.,etal.ThemicrotubuleassociatedproteinAtMAP70-5regulatessecondarywallpatterninginArabidopsiswoodcells[J].CurrentBiology,2010,20:744-749.PesquetE.,ZhangB.,GorzsasA.,etal.Non-cell-autonomouspostmortemlignificationoftrachearyelementsinZinniaelegans[J].ntCell,2013,25:1314-1328.RobischonM.,DuJ.,MiuraE.,etal.ThePopulusclassIIIHDZIP,popREVOLUTA,influencescambiuminitiationandpatterningofwoodystems[J].ntPhysiology,2011,155:1214-1225.SchraderJ.,NilssonJ.,MellerowiczE.,etal.Ahigh-resolutiontranscriptprofileacrossthewood-formingmeristemofporidentifiespotentialregulatorsofcambialstemcellidentity[J].ntCell,2004,16:2278-2292.SinghR.,Ong-AbdullahM.,LowE.T.,etal.Oilpalmgenomesequencerevealsdivergenceofinterfertilespeciesinoldandnewworlds[J].Nature,2013,500:335-SpicerR.,GrooverA..Evolutionofdevelopmentofvascularcambiaandsecondarygrowth[J].NewPhytologist,2010,186:577-592.SuerS.,AgustiJ.,SanchezP.,etal.WOX4impartsauxinresponsivenesstocambiumcellsinArabidopsis[J].ntCell,2011,23:3247-3259.UchidaN.,TasakaM..Regulationofntvascularstemcellsbyendodermis-derivedEPFL-familypeptidehormonesandphloem-expressedERECTA-familyreceptorkinases[J].JournalofExperimentalBotany,2013:ert196.VahalaJ.,FeltenJ.,LoveJ.,etal.Agenome-widescreenforethyleneinducedethyleneresponsefactors(ERFs)inhybridaspenstemidentifiesERFgenesthatmodifystemgrowthandwoodproperties[J].NewPhytologist,2013.200:511-522.WadenbackJ.,vonArnoldS.,EgertsdotterU.,etal.Ligninbiosynthesisintransgen