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[] [] [] [] []75低磷条件下植物根系形态反应及其调控机制_陈磊-第2页
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75低磷条件下植物根系形态反应及其调控机制_陈磊-2
而且在缺磷条件下,蔗糖会显著促进拟南芥突变体pg;[44];;WRKY转录因子家族中WRKY75是拟南芥磷;饥饿反应中被诱导的一个转录因子,WRKY75参与;[96];表现为根毛数量减少,侧根长度和数量降低;植物排根(Clusterroot,又称簇生根)是;2~3mg?g-1[114];[105];;低磷胁迫下,生长素对排根的发生、发育起;着重要作用;[10
而且在缺磷条件下,蔗糖会显著促进拟南芥突变体pgp19、tt4-2及其野生型的侧根数量与长度增加,表明蔗糖参与了缺磷胁迫下生长素对侧根[44]。的发生、发育的调控WRKY转录因子家族中WRKY75是拟南芥磷饥饿反应中被诱导的一个转录因子,WRKY75参与调节磷胁迫下植物根系发育。通过RNA干涉抑制WRKY75基因的表达,植株磷吸收量明显减小,还[96]表现为根毛数量减少,侧根长度和数量降低。植物排根(Clusterroot,又称簇生根)是一种2~3mg?g-1[114]。Ahmad等研究也得到了相似的结果,白羽扇豆排根的形成随着根、茎和叶中磷浓[105]。度的增加而显著下降低磷胁迫下,生长素对排根的发生、发育起着重要作用。低磷胁迫诱导排根启动,外源施用NAA会改变在高磷处理对排根启动的抑制[117]。有试验表明细胞分裂素也参与调控排根发育。根尖产生的细胞分裂素抑制了小侧根的进一步伸[108,109],因此,磷影响排根的长,促进排根形成形成可能与植株体内激素的含量及相对分配有关。Wang等研究表明NO影响缺磷胁迫下白羽扇豆排根的形成。缺磷胁迫增加了白羽扇豆初生根和侧根根尖中NO的产生量,排根中NO的产生量增加更多。使用外源NO合酶和黄嘌呤氧化还原酶的抑制剂可减少缺磷条件下NO产生量,使用外源NO供体硝普钠明显增加白羽扇豆排根[118]。数量5展望特殊的侧根形态结构,侧根上形成象毛刷一样密集而有限生长的一组侧根的根簇。排根起源于中柱鞘[97]细胞,具有严密的时空发育规律。排根在山龙[98]眼科植物中普遍存在,还分布在许多其他植物如豆科、莎草科、桦木科、杨梅科、木麻黄科[99,100]。白羽扇豆(Lupinusalbus)是研究排根形等成的模式植物。缺磷环境下白羽扇豆形成排[100,101],是其适应磷胁迫的一种重要机根。缺磷胁迫下白羽扇豆种子萌发后9d就会形成簇生根,萌发后14d约三分之二的次级根都制是排根[106][102-105]。土壤中磷的不均匀分布影响排根的形[107]低磷胁迫下,植物新陈代谢,生理、生化反应以至根系形态发生适应性改变,而这些形态发育及生理的变化对于增强植物对土壤中磷的吸收,提高磷素利用效率,使植物避免低磷胁迫的伤害具有重要的意义。近十年来,分子生物学技术的不断发展,植物磷吸收利用方面突变体和拟南芥根系生长发育突变体不断被筛选出来,磷素养分调节根系生长发育的信号转导机制的研究取得了很大进展,激素、转录因子、关键蛋白参与调控了低磷胁迫下植物的生长发育(图1),但这些信号之间的相互关系还不甚清楚,信号传递通路中,谁在上游,谁在下游,还缺少研究。结合分子标记、基因定位、转基因、基因融合表达和基因芯片等操作技术,推进缺磷胁迫根系发育功能基因组研究,将有助于深入了解相关的基因表达及相互调控机制。迄今对磷素养分调节根系发育的研究大多是集中在模式植物拟南芥上,有关磷素养分供应影响农作物根系生长、发育的生理与分子机制还需要更深入的研究。借助拟南芥上的研究成果,对作物根系形态的一些性状进行定量测定并进行遗传基础研究,克隆重要根系性状的相关基因,将对培育磷高效优良作物品种具有深远的意义。成与分布,富磷区域内排根明显增多。排根的存在极大地增加根系与土壤的接触面积,增加磷对植物的空间有效性,提高植株对土壤中磷的获取机会。白羽扇豆每厘米基根上有250~500个细根,而同样长度的非排根,每厘米基根上细根数量少于50个[102,108],相对细长的排根可使单位长度基根表面积增加140倍,与土体接触面积增[108]。排根还释放质子、分泌柠檬酸以及加288倍磷酸酶等活化土壤中磷,增加土壤中磷对植物的有[100,109-111]。在缺磷胁迫下白羽扇豆排根分泌的效性有机酸是供磷充足情况下的65倍。其中以柠檬酸为主,占根系分泌有机酸的87%,随着分泌柠檬酸的增加,根际pH值下降,与对照相比,pH值从7.1下降至4.6[109]。排根的发生、发育与植物体内的磷浓度有[100,105,112-114]。叶面喷施磷和分根试验表明,白关羽扇豆的排根形成受植株体内磷含量的调控,而与[99,115,116]。高磷供应下,白外部磷的局部调控无关羽扇豆排根形成的下降与茎和韧皮部组织液中磷浓[112,113]。Li等研究也指出,白羽扇豆排根的度有关形成是受茎中磷浓度的调控,调控的临界浓度是―6―图1低磷胁迫下植物根系形态的调控机制注:实线箭头代表正调控作用,虚线代表负调控作用。空心箭头代表在低磷胁迫条件下铁的升高和植物激素的降低。粗黑框代表转录因子。参考文献:[1]BarberSA,WalkerJM,VaseyEH.Mechanismsforthemovementofplantnutrientsfromthesoilandfertilizertotheplantroot[J].JournalofAgriculturalandFoodChemistry,):204-207.[2]刘建中,李振声,李继云.利用植物自身潜力提高土壤中磷J].生态农业研究,):16-23.的生物有效性[[3][4]M].北京:中国农业出全国土壤普查办公室.中国土壤[版社,1998.鲁如坤,史正元,顾益初.土壤积累态磷研究.II磷肥的表观积累利用率[J].土壤肥料,):286-289.[5]张福锁,崔振岭,王激清,等.中国土壤和植物养分管理.植物学通报,):687现状与改进策略[J]-694.[6]辛景树,徐明岗,田有国,等.耕地质量演变趋势研究―7――――国家级耕地土壤监测数据整编[M].北京:中国农业科学技术出版社,2008.[7]LambersH,ShaneMW,CramerMD,etal.Rootstructureandfunctioningforefficientacquisitionofphosphorus:matchingmorphologicalandphysiologicaltraits[J].AnnalsofBotany,3-713.[8]UnitedStatesGeologicalSurvey.MineralCommoditySumma-ries[R].Washington:UnitedStatesGovernmentPrintingOffice,2009.[9]SteenI.Phosphorusavailabilityinthe21stcentury:Manage-.PhosphorusandPo-mentofanon-renewableresource[J]tassium,:25-31.[10]IsherwoodKF.Mineralfertilizeruseandtheenvironment[M].Paris:InternationalFertilizerIndustryAssociation.U-nitedNationsEnvironmentProgramme,2000.[11]CordellD,DrangertJO,WhiteS.Thestoryofphosphorus:globalfoodsecurityandfoodforthought[J].GlobalEnviron-mentalChange,2-305.[12][13]LambersH,ChapinFS,PonsTL.Plantphysiologicalecology[M].NewYork:Springer,1998.ClarksonDT.Nutrientinterceptionandtransportbyrootsystems[A].In:JohnsonCB(eds).PhysiologicalprocesseslimitingM].London:Butterworths,1981.plantproductivity[[14][15]严小龙,廖红,戈振扬,等.植物根构型特性与磷吸收效J].植物学通报,):511-519.率[ShinR,BergRH,SchachtmanDP.ReactiveoxygenspeciesandroothairsinArabidopsisrootresponsetonitrogen,phos-phorusandpotassiumdeficiency[J].PlantandCellPhysiolo-gy,50-1357.[16]TewariRK,KumarP,SharmaPN.Oxidativestressandan-tioxidantresponsesinyoungleavesofmulberryplantsgrownun-dernitrogen,phosphorusorpotassiumdeficiency[J].JournalofIntegrativePlantBiology,):313-322.[17]HerderGD,VanIG,BeeckmanT,etal.Therootsofanew.TrendsinPlantScience,2010,15greenrevolution[J](11):600-607.[18][19][20]赵华,徐芳森,石磊,等.植物根系形态对低磷胁迫应答.植物学报,):409-417.的研究进展[J].杭李永夫.水稻适应低磷胁迫的营养生理机理研究[D]州:浙江大学,2006.GeorgeTS,FranssonAM,HammondJP,etal.Phosphorusnutritionrhizosphereprocesses,plantresponseandAdaptations[A].BunemannEK,ObersonA,FossardE(eds.),PhosphorusInAction:Biologicalprocessesinsoilphosphoruscycling[M].NewYork:Springer,-271.[21]ShrideviA,Jakkeral,KajjidoniST,etal.Genotypicvaria-tionforroottraitstophosphorusdeficiencyinblackgram(VignamungoL.Hepper)[J].KarnatakaJournalofAgriculturalSci-ences,):946-950.[22]RouachedH,ArpatAB,PoirierY.Regulationofphosphatestarvationresponsesinplants:signalingplayersandcross-[40][39][38][37][36][35][34][33][32][31][30][29][28][27][26][25][24][23]J].MolecularPlant,):288-299.talks[WilliamsonLC,RibriouxS,FitterAH,etal.Phosphatea-vailabilityregulatesrootsystemarchitectureinArabidopsis[J].PlantPhysiology,:875-882.廖红,严小龙.菜豆根构型对低磷胁迫的适应性变化及基J].植物学报,):158-163.因型差异[LiaoH,YanXL,RubioG,etal.Geneticmappingofbasalrootgravitropismandphosphorusacquisitionefficiencyincommonbean[J].FunctionalPlantBiology,9-970.BatesTR,LynchJP.Roothairsconferacompetitiveadvan-.PlantandSoil,tageunderlowphosphorusavailability[J]:243-250.刘灵,廖红,王秀荣,等.不同根构型大豆对低磷的适应性变化及其与磷效率的关系[J].中国农业科学,):.廖红,戈振扬,严小龙.水磷耦合胁迫下植物磷吸收的理.科学通报,):想根构型:模拟与应用[J]641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