调节转运蛋白AtNRT1.1、AtNRT2.1及PdAMT1.1基因表达对拟南芥和杨树氮素吸收的影响研究
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摘要
氮素营养是调节植物生长发育的主要外界环境因子之一。而硝酸盐(NO3-)和铵盐(NH4+)是植物根部从土壤中吸收和利用氮素的两种主要的稳定形态的氮源。目前研究证明,植物对NH4+、N03的吸收利用受氮素转运蛋白家族中的多个基因调控。其中,NRT1.1、NRT2.1是NO3-转运系统中的关键基因,而AMT1.1基因是NH4+转运系统中的关键基因。通过调节这些关键转运基因的表达可以提高植物氮素吸收效率。
本研究分别以乙烯调节、生长素调节及转基因技术这三种手段来调节氮素转运蛋白AtNRT2.1、AtNRT1.1和PdAMT1.1基因的表达水平,从而来影响拟南芥和杨树的氮素营养吸收效率。
在乙烯调节方面,作者运用气相色谱法、扫描离子选择电极法(SIET)、实时荧光定量法(qRT-PCR)、GUS染色法等技术来检测拟南芥AtNRT2.1基因在NO3-吸收途径与乙烯合成及信号转导通路的互作网络中的地位。结果发现：拟南芥(Col-0)植株在低NO3-浓度(LN)处理下乙烯合成量有显著增加,并且乙烯信号通路中的关键因子CTR1、 EIN3和EIL1基因表达量受到调节,在乙烯报告基因株系Col-0/EBS:GUS、 ein3-leil1-1/EBS:GUS和ctr1-1/EBS:GUS中的EBS:GUS活性均有显著增加。LN处理可以诱导AtNRT2.1基因表达,从而增加高亲和的NO3-吸收速率。通过对比NO3-转运蛋白突变体nrt1.1、nrt2.1与Col-0株系中的乙烯合成量及EBS:GUS活性差异,证实了LN处理诱导的AtNRT2.1基因上调可以促进乙烯合成、增强乙烯信号通路强度；同时,乙烯又可负调控AtNRT2.1基因表达,降低高亲和的NO3-吸收速率。综上所述,我们揭示了在LN处理下,AtNRT2.1基因表达与乙烯合成及信号转导途径间存在一个负反馈调节环,这样在外界NO3-缺乏胁迫下的NO3-吸收就可以达到一个内部相对平衡的状态。
在生长素调节方面,为研究生长素信号对植物NO3-营养吸收的调控作用,作者采用SIET技术和qRT-PCR技术,测定拟南芥野生型Col-0、生长素过表达突变体yuc1-D以及生长素通路缺失突变体axr1-12三种株系初始根中NO3-吸收速率以及AtNRT1.1基因表达量的差异；并进一步检测Col-0株系以及NO3-转运蛋白突变体nrt1.1株系在正常条件(CK)或施加外源IAA及生长素极性运输抑制剂2,3,5-三碘苯甲酸(TIBA)处理下的NO3-流速和AtNRT1.1基因表达量的差异。结果表明yuc1-D株系的N03-吸收速率以及AtNRT1.1基因表达量相比Col-0株系均有大幅增加,而axr1=12株系的N03-吸收速率以及AtNRT1.1基因表达量相比Col-0株系显著降低；在Col-0株系中施加外源IAA对N03-吸收速率以及AtNRT1.1基因表达量有明显促进作用,而施加TIBA的效果反之,说明生长素对硝酸盐吸收有增强效应。而nrtl.1株系在CK、IAA.TIBA处理下N03-吸收速率差异较Col-0株系不明显,揭示了AtNRT1.1基因在生长素促进N03-吸收的调控途径中所起的重要作用。
在转基因方面,作者从欧美杨NE-19中筛选并克隆了高亲和NH4+转运蛋白家族中的关键基因PdAMT1.1。经测序及蛋白结构分析发现,该基因片段大小为1,495bp,共编码498个氨基酸,包含9个跨膜蛋白结构域和1个保守结构域。组织特异性分析表明PdAMT1.1在根中的表达量要明显高于于芽、叶及茎中的表达量。通过亚细胞定位分析表明其定位于膜系统中。在模式植物拟南芥中过表达PdAMT1.1基因,发现在氮缺乏条件下,转基因株系OxPdAMT1.1的叶而积、株高等生理表型及NH4+吸收效率要明显高于其它株系,突变体amt1.1的表型及NH4+吸收效率最弱,而突变体回补株系amt1.1/PdAMT1.1的表型及NH4+吸收功能基本与Col-0及转空载体Col-O/V株系持平。从而验证了PdAMT1.1基因在促进植物生长、提高NH4+吸收效率方面发挥重要作用。然后我们通过叶盘侵染法进一步将PdAMT1.1基因转化进三倍体毛白杨中。通过氯酚红法检测、GUS检测、PCR检测等一系列手段,验证已获得6个转PdAMT1.1基因株系及3个转空载体株系。对缺氮处理下三倍体毛白杨的野生型WT、转空载体WT/V及过表达株系OxPdAMT1.1进行表型和功能分析,结果证明过表达株系OxPdAMT1.1表型及NH4+吸收效率均明显优于WT和WT/V株系。从而验证了通过转基因手段使得PdAMT1.1基因超量表达,可以获得能高效吸收利用氮素营养的杨树新品种。
Nitrogen availability is a major environmental factor that regulates plant growth and development. Nitrate (NO3-) and ammonium (NH4+) represent the most readily available forms of nitrogen for root absorption from the soil. Based on recent studies, there are many genes from different nitrogen transporter families responsible for NH4+and NO3-absorption. NRT1.1and NRT2.1are two crucial genes in NO3-transport system, and AMT1.1is a key gene in NH4+transport system. Regulating the expression of these genes could improve the efficiency of nitrogen absorption in plants.
We regulated the expression of nitrogen transporter genes AtNRT2.1, AtNRT1.1and PdAMT1.1by ethylene mediation, auxin mediation and transgenic technology, respectively, to improve the efficiency of nitrogen absorption in Arabidopsis and poplar.
Gas chromatography, scanning ion-selective electrode (SIET), quantitative real-time PCR (qRT-PCR) and GUS reporter assay techniques were used to explore the molecular interaction between AtNRT2.1transcript levels and the ethylene signaling pathway under nitrate deficiency. We reported a low nitrate (LN) treatment-induced rapid burst of ethylene production and regulated expression of ethylene signalling components CTR1, EIN3and EIL1in wild-type Arabidopsis thaliana (Col-0) seedlings, and enhanced ethylene response reporter EBS:GUS activity in both Col-0and the ethylene mutants ein3-leill-1and ctr1-1. LN treatment also caused up-regulation of AtNRT2.1expression, which was responsible for an enhanced high-affinity nitrate uptake. Comparison of ethylene production and EBS:GUS activity between mi1.1, nrt2.1mutants and Col-0indicated that this up-regulation of AtNRT2.1expression caused a positive effect on ethylene biosynthesis and signaling under LN treatment. On the other hand, ethylene downregulated AtNRT2.1expression and reduced the high-affinity nitrate uptake. Together, these findings uncover a negative feedback loop between AtNRT2.1expression and ethylene biosynthesis and signalling under nitrate deficiency, which may contribute to finely tuning of plant nitrate acquisition during exploring dynamic soil conditions.
In order to study the modulating effect of auxin signaling on nitrate uptake in plants, SIET and qRT-PCR techniques were used to determine the net plasma membrane NO3-fluxes and the expression of nitrate transporter AtNRT1.1in primary roots of wild-type Arabidopsis Col-0, auxin-overproducing mutant yucl-D, and auxin-insensitive mutant axrl-12. Moreover, we examined the net plasma membrane NO3-fluxes and the expression of AtNRT1.1in primary roots of Col-0and dual-affinity nitrate transporter mutant nrtl.l seedlings under the normal condition (CK), the treatment with exogenous IAA, or the treatment with the auxin efflux inhibitor2.3.5-triidobenzoid acid (TIBA). The nitrate uptake and the transcript level of AtNRT1.1in yucl-D seedlings were remarkably enhanced, whereas those in axrl-12seedlings declined compared with Col-0seedlings. The exogenous IAA treatment enhanced the nitrate uptake and the transcript level of AtNRT1.1in Col-0seedlings, while the TIBA treatment had an opposite effect. Furthermore, the differences of the nitrate uptake and the expression of AtNRT1.1in nrt1.1seedlings among CK, IAA and TIBA treatments were insignificant compared with Col-0seedlings, revealing the important role that AtNRT1.1gene played in nitrate uptake pathway response to auxin.
We cloned a key gene for the high-affinity NH4+uptake, PdAMT1.1, from Populus nigra×(Populus deltoides×Populus nigra). After genetic sequencing and protein structure analysis, we found the PdAMT1.1cDNA is1,495bp in length and encodes498amino acids. PdAMT1.1protein encodes9transmembrane domains and one conserved domain. The tissue-specific expression analysis indicates PdAMT1.1was expressed more highly in roots than in stems, buds and leaves. The subcellular localization analysis indicates PdAMT1.1localized in membrane system. Transgenic Arabidopsis plants that constitutively expressed the PdAMT1.1gene were constructed. We found under nitrogen deficiency conditions, the physiological phenotypes such as leaf areas and plant height, and the NH4+influxes in transgenic OxPdAMT1.1seedlings were much better than other seedlings, while those in amtl.l seedlings were most inferior. The complemented line amt1.1/PdAMT1.1demonstrated similar phenotype and NH4+uptake efficiency as wild type Col-0seedlings and the empty vector transgenic line Col-O/V. These results suggest that PdAMTI.l gene plays an important role in plant growth and NH4+absorption. Then we transformed PdAMT1.1gene into triploid white poplar by leaf disk method. Six transgenic OxPdAMT1.1lines and three empty vector transgenic lines were checked by Chlorophenol-Red assay, GUS assay and PCR analysis. To study the biological functions of triploid white poplar wild type seedlings WT, the empty vector transgenic line WT/V and transgenic OxPdAMT1.1seedlings, we found OxPdAMT1.1seedlings have much better phenotypes and NH4+uptake efficiency than WT and WT/V seedlings. Thus we proved the overexpression of PdAMT1.1gene by transgenic technology could generate new poplar species which have high nitrogen absorption efficiency.
本研究分别以乙烯调节、生长素调节及转基因技术这三种手段来调节氮素转运蛋白AtNRT2.1、AtNRT1.1和PdAMT1.1基因的表达水平,从而来影响拟南芥和杨树的氮素营养吸收效率。
在乙烯调节方面,作者运用气相色谱法、扫描离子选择电极法(SIET)、实时荧光定量法(qRT-PCR)、GUS染色法等技术来检测拟南芥AtNRT2.1基因在NO3-吸收途径与乙烯合成及信号转导通路的互作网络中的地位。结果发现：拟南芥(Col-0)植株在低NO3-浓度(LN)处理下乙烯合成量有显著增加,并且乙烯信号通路中的关键因子CTR1、 EIN3和EIL1基因表达量受到调节,在乙烯报告基因株系Col-0/EBS:GUS、 ein3-leil1-1/EBS:GUS和ctr1-1/EBS:GUS中的EBS:GUS活性均有显著增加。LN处理可以诱导AtNRT2.1基因表达,从而增加高亲和的NO3-吸收速率。通过对比NO3-转运蛋白突变体nrt1.1、nrt2.1与Col-0株系中的乙烯合成量及EBS:GUS活性差异,证实了LN处理诱导的AtNRT2.1基因上调可以促进乙烯合成、增强乙烯信号通路强度；同时,乙烯又可负调控AtNRT2.1基因表达,降低高亲和的NO3-吸收速率。综上所述,我们揭示了在LN处理下,AtNRT2.1基因表达与乙烯合成及信号转导途径间存在一个负反馈调节环,这样在外界NO3-缺乏胁迫下的NO3-吸收就可以达到一个内部相对平衡的状态。
在生长素调节方面,为研究生长素信号对植物NO3-营养吸收的调控作用,作者采用SIET技术和qRT-PCR技术,测定拟南芥野生型Col-0、生长素过表达突变体yuc1-D以及生长素通路缺失突变体axr1-12三种株系初始根中NO3-吸收速率以及AtNRT1.1基因表达量的差异；并进一步检测Col-0株系以及NO3-转运蛋白突变体nrt1.1株系在正常条件(CK)或施加外源IAA及生长素极性运输抑制剂2,3,5-三碘苯甲酸(TIBA)处理下的NO3-流速和AtNRT1.1基因表达量的差异。结果表明yuc1-D株系的N03-吸收速率以及AtNRT1.1基因表达量相比Col-0株系均有大幅增加,而axr1=12株系的N03-吸收速率以及AtNRT1.1基因表达量相比Col-0株系显著降低；在Col-0株系中施加外源IAA对N03-吸收速率以及AtNRT1.1基因表达量有明显促进作用,而施加TIBA的效果反之,说明生长素对硝酸盐吸收有增强效应。而nrtl.1株系在CK、IAA.TIBA处理下N03-吸收速率差异较Col-0株系不明显,揭示了AtNRT1.1基因在生长素促进N03-吸收的调控途径中所起的重要作用。
在转基因方面,作者从欧美杨NE-19中筛选并克隆了高亲和NH4+转运蛋白家族中的关键基因PdAMT1.1。经测序及蛋白结构分析发现,该基因片段大小为1,495bp,共编码498个氨基酸,包含9个跨膜蛋白结构域和1个保守结构域。组织特异性分析表明PdAMT1.1在根中的表达量要明显高于于芽、叶及茎中的表达量。通过亚细胞定位分析表明其定位于膜系统中。在模式植物拟南芥中过表达PdAMT1.1基因,发现在氮缺乏条件下,转基因株系OxPdAMT1.1的叶而积、株高等生理表型及NH4+吸收效率要明显高于其它株系,突变体amt1.1的表型及NH4+吸收效率最弱,而突变体回补株系amt1.1/PdAMT1.1的表型及NH4+吸收功能基本与Col-0及转空载体Col-O/V株系持平。从而验证了PdAMT1.1基因在促进植物生长、提高NH4+吸收效率方面发挥重要作用。然后我们通过叶盘侵染法进一步将PdAMT1.1基因转化进三倍体毛白杨中。通过氯酚红法检测、GUS检测、PCR检测等一系列手段,验证已获得6个转PdAMT1.1基因株系及3个转空载体株系。对缺氮处理下三倍体毛白杨的野生型WT、转空载体WT/V及过表达株系OxPdAMT1.1进行表型和功能分析,结果证明过表达株系OxPdAMT1.1表型及NH4+吸收效率均明显优于WT和WT/V株系。从而验证了通过转基因手段使得PdAMT1.1基因超量表达,可以获得能高效吸收利用氮素营养的杨树新品种。
Nitrogen availability is a major environmental factor that regulates plant growth and development. Nitrate (NO3-) and ammonium (NH4+) represent the most readily available forms of nitrogen for root absorption from the soil. Based on recent studies, there are many genes from different nitrogen transporter families responsible for NH4+and NO3-absorption. NRT1.1and NRT2.1are two crucial genes in NO3-transport system, and AMT1.1is a key gene in NH4+transport system. Regulating the expression of these genes could improve the efficiency of nitrogen absorption in plants.
We regulated the expression of nitrogen transporter genes AtNRT2.1, AtNRT1.1and PdAMT1.1by ethylene mediation, auxin mediation and transgenic technology, respectively, to improve the efficiency of nitrogen absorption in Arabidopsis and poplar.
Gas chromatography, scanning ion-selective electrode (SIET), quantitative real-time PCR (qRT-PCR) and GUS reporter assay techniques were used to explore the molecular interaction between AtNRT2.1transcript levels and the ethylene signaling pathway under nitrate deficiency. We reported a low nitrate (LN) treatment-induced rapid burst of ethylene production and regulated expression of ethylene signalling components CTR1, EIN3and EIL1in wild-type Arabidopsis thaliana (Col-0) seedlings, and enhanced ethylene response reporter EBS:GUS activity in both Col-0and the ethylene mutants ein3-leill-1and ctr1-1. LN treatment also caused up-regulation of AtNRT2.1expression, which was responsible for an enhanced high-affinity nitrate uptake. Comparison of ethylene production and EBS:GUS activity between mi1.1, nrt2.1mutants and Col-0indicated that this up-regulation of AtNRT2.1expression caused a positive effect on ethylene biosynthesis and signaling under LN treatment. On the other hand, ethylene downregulated AtNRT2.1expression and reduced the high-affinity nitrate uptake. Together, these findings uncover a negative feedback loop between AtNRT2.1expression and ethylene biosynthesis and signalling under nitrate deficiency, which may contribute to finely tuning of plant nitrate acquisition during exploring dynamic soil conditions.
In order to study the modulating effect of auxin signaling on nitrate uptake in plants, SIET and qRT-PCR techniques were used to determine the net plasma membrane NO3-fluxes and the expression of nitrate transporter AtNRT1.1in primary roots of wild-type Arabidopsis Col-0, auxin-overproducing mutant yucl-D, and auxin-insensitive mutant axrl-12. Moreover, we examined the net plasma membrane NO3-fluxes and the expression of AtNRT1.1in primary roots of Col-0and dual-affinity nitrate transporter mutant nrtl.l seedlings under the normal condition (CK), the treatment with exogenous IAA, or the treatment with the auxin efflux inhibitor2.3.5-triidobenzoid acid (TIBA). The nitrate uptake and the transcript level of AtNRT1.1in yucl-D seedlings were remarkably enhanced, whereas those in axrl-12seedlings declined compared with Col-0seedlings. The exogenous IAA treatment enhanced the nitrate uptake and the transcript level of AtNRT1.1in Col-0seedlings, while the TIBA treatment had an opposite effect. Furthermore, the differences of the nitrate uptake and the expression of AtNRT1.1in nrt1.1seedlings among CK, IAA and TIBA treatments were insignificant compared with Col-0seedlings, revealing the important role that AtNRT1.1gene played in nitrate uptake pathway response to auxin.
We cloned a key gene for the high-affinity NH4+uptake, PdAMT1.1, from Populus nigra×(Populus deltoides×Populus nigra). After genetic sequencing and protein structure analysis, we found the PdAMT1.1cDNA is1,495bp in length and encodes498amino acids. PdAMT1.1protein encodes9transmembrane domains and one conserved domain. The tissue-specific expression analysis indicates PdAMT1.1was expressed more highly in roots than in stems, buds and leaves. The subcellular localization analysis indicates PdAMT1.1localized in membrane system. Transgenic Arabidopsis plants that constitutively expressed the PdAMT1.1gene were constructed. We found under nitrogen deficiency conditions, the physiological phenotypes such as leaf areas and plant height, and the NH4+influxes in transgenic OxPdAMT1.1seedlings were much better than other seedlings, while those in amtl.l seedlings were most inferior. The complemented line amt1.1/PdAMT1.1demonstrated similar phenotype and NH4+uptake efficiency as wild type Col-0seedlings and the empty vector transgenic line Col-O/V. These results suggest that PdAMTI.l gene plays an important role in plant growth and NH4+absorption. Then we transformed PdAMT1.1gene into triploid white poplar by leaf disk method. Six transgenic OxPdAMT1.1lines and three empty vector transgenic lines were checked by Chlorophenol-Red assay, GUS assay and PCR analysis. To study the biological functions of triploid white poplar wild type seedlings WT, the empty vector transgenic line WT/V and transgenic OxPdAMT1.1seedlings, we found OxPdAMT1.1seedlings have much better phenotypes and NH4+uptake efficiency than WT and WT/V seedlings. Thus we proved the overexpression of PdAMT1.1gene by transgenic technology could generate new poplar species which have high nitrogen absorption efficiency.
引文
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