富含亮氨酸的类受体蛋白激酶在南极丝瓜藓（Pohlia nutans）适应极端环境中的作用

发布时间：2018-05-26 21:40

本文选题：转录组测序 + 富含亮氨酸的类受体蛋白激酶　；参考：《山东大学》2017年博士论文

【摘要】：南极地区由陆地和海洋组成,是地球上的极寒、极冰、极风和极干的区域。南极独特的地理构成和极端的气候使得南极沿海地区隐花植物(苔藓和地衣)的种类和特征有别于世界其他地区的植物资源,具有独特的抗低温、耐高盐、耐干旱、抗辐射等适应能力。苔藓是南极大陆主要的植物类群,其逆境适应机制仍不清楚。富含亮氨酸的类受体蛋白激酶(Leucine-rich repeat receptor-like kinases,LRR-RLK)作为最初感知外界环境变化和传导信号类受体蛋白激酶最大的亚家族,可能在苔藓适应极地严酷环境过程中发挥着关键性作用。本研究中,我们利用Illumina公司的高通量测序技术分析了南极丝瓜藓(Pohlia nutans)在高盐胁迫下的转录组,从基因组水平上筛选了LRR-RLK 亚家族基因并分析了其对盐胁迫的响应特征;筛选出3个对盐胁迫正响应的LRR-RLKs(PnLRR-RLK19、PnLRR-RLK27和PnLRR-RLK44)进行了深入研究,探讨了其在模式植物拟南芥和/或小立碗藓中的功能,并探讨了其可能的分子机制;对于揭示南极苔藓适应极端环境的作用机制,促进极地独特基因资源的开发和利用具有一定的理论意义和应用价值。1植物PnLRR-RLK基因的转录组分析利用高通量测序技术分析了南极丝瓜藓(Pohlia nuans)在正常条件和高盐胁迫下(200 mM NaCl,1 h)的基因转录组特征。转录组测序后最终共得到72922个单一基因序列,平均长度为704 bp;单一基因序列大于500 bp的共有23747个。以转录组数据库为基础,在南极丝瓜藓筛选出56个候选的富含亮氨酸类受体蛋白激酶的蛋白序列,命名为PnLRR-RLK1-56,根据胞内激酶结构域将LRR-RLK分为11个亚家族(LRR Ⅰ-ⅩⅢ)。盐胁迫下有4个PnLRR-RLKs表达水平显著上调,有10个PnLRR-RLKs的表达水平显著下调,进一步研究了在盐胁迫条件下3个正响应PnLRR-RLKs(19/27/44)的生物信息学分析,发现3个基因在高盐、低温、干旱、紫外辐射和脱落酸(ABA)处理下显著上调表达。蛋白序列比对分析表明,PnLRR-RLK19/27/44具有保守的蛋白激酶结构域,与其他物种PnLRR-RLKs的同源性为 41%-60%;PnLRR-RLK19 与 PnLRR-RLK27 和 PnLRR-RLK44 的同源性分别为43.7%、16.9%和19.6%。系统进化树分析发现,南极苔藓基因PnLRR-RLK19 与 PnLRR-RLK27 属于 LRRⅡ型的 RLK 亚家族,PnLRR-RLK44属于LRR Ⅺ型的RLK亚家族。我们进一步对3个受多种胁迫诱导的PnLRR-RLKs基因进行深入的生物学功能研究。1.1 PnLRR-RLK27提高了植株对盐胁迫和氧化胁迫的抗性将GFP和PnLRR-RLK27:GFP的瞬时表达载体分别转化拟南芥和小立碗藓原生质体,暗培养一段时间后,共聚焦显微镜观察荧光分布发现,对照组GFP绿色荧光均匀分布在细胞膜、细胞质和细胞核中;PnLRR-RLK27:GFP绿色荧光可能在细胞膜上分布。进一步利用与H+-ATPase-RFP载体(细胞膜定位的marker基因,发红色荧光)共转或细胞膜特异性染料FM4-64(发红色荧光)分析PnLRR-RLK27的亚细胞定位,结果发现PnLRR-RLK27:GFP绿色荧光与H+-ATPase-RFP在细胞膜上红色荧光或FM4-64的红色荧光完全重叠,呈黄色;表明,PnLRR-RLK27定位在细胞膜。在小立碗藓中的功能及机制研究:通过同源重组构建小立碗藓过表达载体pTFH15.3:PnLRR-RLK27,通过遗传转化使PnLRR-RLK27在小立碗藓中组成型表达。取单个长度为3 mm野生型和过表达PnLRR-RLK27小立碗藓植株的茎尖放入含0 mM和125 mM NaCl的BCD中,培养7周。发现正常情况下,PnLRR-RLK27不影响小立碗藓的生长(直径均为13 mm左右);而盐处理下(125 mM BCD),过表达PnLRR-RLK27显著提高了植株对盐胁迫的抗性,野生型植株配子体直径为4.7mm,过表达PnLRR-RLK27植株配子体为10.0mm左右。在含10μM和15μM ABA的BCD培养基上,过表达PnLRR-RLK27植株配子体直径比野生型大1.6倍和1.8倍。与野生型植株相比,盐胁迫下转基因小立碗藓的盐胁迫响应基因(PpENA2、PpSHP1和PpSHP2)、和胁迫诱导基因(PpDBF、PpCOR47和PpCORTMC-AP3)和ABA信号途径负调控基因(PpABI3a和PpABI3b)的表达量明显提高。表明,过表达PnLRR-RLK27显著增强了小立碗藓在配子体生长阶段对盐胁迫抗性和降低了对ABA的敏感性。在拟南芥中的功能及机制研究:构建了PnLRR RLK27的pROK2载体,通过农杆菌侵染拟南芥花序,获得了过表达PnLRR-RLK27转基因拟南芥纯系植株(以下简称转基因植株)。盐胁迫下,转基因植株种子萌发率比野生型植株高1.5倍,转基因植株主根是野生型植株1.3倍左右。ABA处理下,转基因植株的萌发率是野生型植株的2倍,主根比野生型植株长10.0倍。在H202处理下,转基因植株主根比野生型长1.4倍,侧根数多1.6倍。表明,过表达PnLRR-RLK27提高了拟南芥对盐胁迫和氧化胁迫的抗性,降低了对ABA的敏感性。与野生型拟南芥相比,盐处理下过表达PnLRR-RLK27降低了植株H202和MDA的含量,增加了脯氨酸的含量,提高了抗氧化酶超氧化物岐化酶(superoxide dismutase,SOD)、过氧化物酶(peroxidase,POD)和过氧化氢酶(catalase,CAT)的活性,上调了抗氧化酶基因At4PX1、AtAPX2、AtCAT2和AtZAT10的表达水平。盐处理下,盐胁迫响应基因(AtHKT和AtSOS3)和ABA信号途径基因(AtABF3、AtMYB2、AtRD22、AtDREB2A、AtRD29A、AtRD29B、AtKIN1 和 AtCOR47)的表达量在转基因植株中比野生型明显提高。以上结果表明,PnLRR-RLK27基因的功能保守,在两种模式植物中功能类似,均能显著提高植物对盐胁迫抗性,其作用机制可能与提高植物对ROS的清除能力以及胁迫相关激素ABA信号途径相关。1.2 PnLRR-RLK19提高了植株对盐胁迫和干旱胁迫的抗性亚细胞定位分析发现,PnLRR-RLK19在细胞膜分布。PnLRR-RLK19在拟南芥中过表达,对植物的正常生长没有影响。盐处理后,转基因植株种子萌发率比野生型植株高25.0%,转基因植株的主根比野生型植株长2.0倍。在0.5 μMABA处理下,转基因植物种子萌发率比野生型植株高11.5%以上;在7.5 μM ABA处理后,转基因植株的主根比野生型植株长1.5倍。H202和甘露醇处理后,过表达植株侧根数比野生型植株分别多1.5倍和2.0-4.0倍。盆栽实验显示,盐处理后,转基因植株叶片黄化比野生型拟南芥少;干旱处理后,转基因植株的失水率较低。盐胁迫处理后,转基因植株ABA信号途径基因(AtAREB1、AtAREB2、AtABF3、AtMYB2、AtDREB2A、AtRD22和AtRD29A)和盐胁迫响应基因(AtP5CS1和AtSOS3)均显著上调表达。干旱处理后,转基因植株ABA信号途径基因(AtABF3、AtMYB2、AtDREB2A、AtRD22、AtRD29A、AtRD29和AtKIN1)显示明显上调的表达模式。以上结果表明,PnLRR-RLK19不仅能够提高了植物对盐胁迫的抗性,也增强了植物对干旱胁迫及氧化胁迫的抗性;其机制与盐胁迫信号途径和ABA信号途径正相关。1.3 PnLRR-RLK44提高了植株对盐胁迫和脱落酸的抗性PnLRR-RLK44定位于细胞膜。构建了PnLRR-RLK4 的过表达载体,利用农杆菌侵染拟南芥,筛选和繁种,最终获得过量表达的转基因纯合植株。在100 mMNaCl培养基上,过表达植株的萌发率比野生型植株高17.6%,转基因植株主根比野生型长28.1%。在0.5 μMABA处理下,转基因植株的种子萌发率比野生型植株高20.0%,主根比野生型长28.9%。进而,利用Real-time qPCR技术检测了盐胁迫信号途径和ABA途径中相关的胁迫响应基因的表达情况,发现过表达植株盐胁迫离子通道蛋白基因(AtHKT1和AtSOS3)、盐胁迫响应基因(AtP5CS1和AtADH1)和和ABA不敏感基因ABI1都呈显著的上调表达。同时,ABA合成途径关键基因(AtNCED3、AtABA1 和AtAAO3)和ABA早期响应基因(AtMYB2、AtRD22、AtRD29A和AtDREB2A)均下调表达。这些结果表明,PnLRR-RLK44通过参与盐胁迫信号途径和与ABA信号途径的负调控因子作用来提高植物的耐盐能力。综上所述,PnLRR-RLK27提高了植物对盐胁迫抗性和降低了对ABA的敏感性,其作用机制可能与提高植物对ROS的清除能力以及盐胁迫/ABA信号途径相关。PnLRR-RLK19不仅提高了植物对盐胁迫抗性和ABA不敏感性,也增强了植物对干旱胁迫及氧化胁迫的抗性,盐胁迫、渗透胁迫和ABA信号途径相关基因表达上调。PnLRR-RLK44通过参与盐胁迫信号途径和与ABA信号途径的负调控因子作用来提高植物的耐盐能力,ABA合成和响应途径下调,ABA负调控途径上调。PnLRR-RLK09/27/44同属一个基因亚家族,其基因功能有类似之处,但也有一定的分化,在提高植物的抗逆机制中有自己独特之处。2 一个泛素连接酶在南极苔藓对非生物胁迫应答中的作用E3泛素连接酶(E3 ubiquitin ligase,E3)可以特异地识别靶蛋白,在植物生长发育和对非生物胁迫响应的过程中发挥着重要的作用。但是,泛素连接酶在低等植物中作用的研究目前还没有报道。本文从南极苔藓中克隆了一个泛素连接酶基因,命名为PnE3。PnE3基因的cDNA含有1个2097 bp开放性阅读框,编码698个氨基酸,分子量为76.2 kDa,等电点为6.79。氨基酸序列分析发现,PnE3包含1个Ubox结构域(氨基酸288-351)、4个ARM重复结构域(氨基酸418-625)。低温、高盐、干旱和ABA处理均能诱导该基因表达上调,表明PnE3基因参与南极苔藓对非生物胁迫响应。将PnE3:GFP的瞬时表达载体转化拟南芥,发现,PnE3:GFP绿色荧光主要在细胞质中分布,表明,PnE3定位在细胞质。在小立碗藓中的功能及机制研究:125 mM NaCl处理下,野生型植株配子体直径比转基因植株大1.4倍;5 μM ABA处理下,野生型植株配子体直径比转基因植株大1.6倍。ABA信号途径基因PpAB13A、PpABI3B和PpABI3C在转基因小立碗藓中均下调表达。在拟南芥中的功能及机制研究:盐处理下,野生型植株的子叶张开率比过表达系高40%,野生型植株的主根根长比过表达系长2.4倍。ABA处理下,野生型植株的子叶张开率比过表达系高25.6%,野生型植株的主根比过表达系长3.7倍。盐胁迫下,胁迫响应基因AtABI3、AtABFA、AtDREB2A和AtRD29A在转基因拟南芥的表达水平显著下调。综上所述,PnE3基因功能保守,在两种模式植物中均能提高植物对盐和ABA的敏感性,其作用机制主要与泛素化修饰植物ABA途径响应基因相关。
[Abstract]:The Antarctic region is composed of land and sea. It is the extreme cold, polar ice, extreme wind and extreme dry areas on the earth. The unique geographical composition and extreme climate of the Antarctic make the species and characteristics of the Antarctic Coastal Areas (moss and lichen) different from the plant resources in other parts of the world, and have unique resistance to low temperature, salt tolerance, drought resistance and resistance. Bryophyte is a major plant group in the Antarctic continent, and its adaptation mechanism is still unclear. Leucine-rich repeat receptor-like kinases (LRR-RLK), which is rich in leucine, is the first subfamily to perceive the changes in the external environment and the largest protein kinase of conduction signals, and may be suitable for mosses. In this study, we should play a key role in the extreme environmental process. In this study, we used Illumina company's high throughput sequencing technology to analyze the transcriptional group of Pohlia nutans under high salt stress. We screened the LRR-RLK subfamily gene from the genome level and analyzed its response to salt stress; 3 of them were screened out. LRR-RLKs (PnLRR-RLK19, PnLRR-RLK27 and PnLRR-RLK44) of positive response to salt stress were studied in depth, and their functions in the model plant Arabidopsis and / or small erect moss were discussed, and their possible molecular mechanisms were discussed. The mechanism of revealing the adaptation of Antarctic moss to the extreme environment and the development of the unique genetic resources of polar regions were also discussed. The transcriptome analysis of the PnLRR-RLK gene of.1 plants with a certain theoretical and applied value was used to analyze the gene transcriptome characteristics of the Antarctic silk moss moss (Pohlia nuans) under normal conditions and high salt stress (200 mM NaCl, 1 h). After the transcriptional group was sequenced, a total of 72922 single gene sequences were obtained and the average length was long. A total of 704 BP; a total of 23747 single gene sequences greater than 500 BP. Based on the transcriptional database, 56 candidate protein kinases rich in leucine receptor protein kinase were screened in the Antarctic gourd moss, named PnLRR-RLK1-56, and LRR-RLK was divided into 11 subfamilies (LRR I - III) based on the intracellular kinase domain. 4 under Salt Stress The expression level of PnLRR-RLKs was significantly up-regulated and the expression level of 10 PnLRR-RLKs decreased significantly. Further study of bioinformatics analysis of 3 positive responses to PnLRR-RLKs (19/27/44) under salt stress showed that 3 genes were significantly up-regulated in high salt, low temperature, drought, ultraviolet radiation and decuquic acid (ABA) treatment. The analysis shows that PnLRR-RLK19/27/44 has a conservative protein kinase domain, and the homology of PnLRR-RLKs with other species is 41%-60%. The homology of PnLRR-RLK19 with PnLRR-RLK27 and PnLRR-RLK44 is 43.7%. 16.9% and 19.6%. phylogenetic tree analysis found that the Antarctic moss gene PnLRR-RLK19 and PnLRR-RLK27 belong to RLK subtype of LRR II. Family, PnLRR-RLK44 belongs to the LRR RLK subfamily. We further study the biological function of 3 PnLRR-RLKs genes induced by multiple stresses..1.1 PnLRR-RLK27 enhanced the plant resistance to salt stress and oxidative stress. The transient expression vectors of GFP and PnLRR-RLK27:GFP were transformed into Arabidopsis and small erect bowl moss, respectively. After a period of dark culture, the fluorescence distribution of the GFP was observed by confocal microscopy. The green fluorescence of the control group was distributed evenly in the cell membrane, cytoplasm and nucleus, and the green fluorescence of PnLRR-RLK27:GFP might be distributed on the cell membrane. Further use of the H+-ATPase-RFP vector (the marker gene located in the cell membrane, the red fluorescence) was further used. Or cell membrane specific dyestuff FM4-64 (red fluorescence) analysis of the subcellular localization of PnLRR-RLK27, the results showed that the PnLRR-RLK27:GFP green fluorescence overlapped the red fluorescence of the cell membrane or the red fluorescence of FM4-64 in the cell membrane. It showed that PnLRR-RLK27 was located in the cell membrane. The function and mechanism in the small erect moss A small bowl of moss overexpression vector pTFH15.3:PnLRR-RLK27 was constructed by homologous recombination, and PnLRR-RLK27 was formed in the small bowl of mosses by genetic transformation. A single length of 3 mm wild type and over expressed PnLRR-RLK27 small erect moss plants were placed in BCD containing 0 mM and 125 mM NaCl, and cultured for 7 weeks. There was no influence on the growth of small erect moss (about 13 mm in diameter), while under salt treatment (125 mM BCD), overexpression of PnLRR-RLK27 significantly increased the resistance to salt stress. The gametophyte diameter of the wild type plant was 4.7MM, and the overexpressed PnLRR-RLK27 plant gametophyte was about 10.0mm. On the BCD medium containing 10 mu M and 15 u M ABA, the overexpressed PnLRR-RLK27 The plant gametophyte diameter was 1.6 times larger and 1.8 times larger than that of the wild type. Compared with the wild type, the salt stress response genes (PpENA2, PpSHP1 and PpSHP2), and the negative regulatory genes (PpABI3a and PpABI3b) of the stress induced genes (PpDBF, PpCOR47 and PpCORTMC-AP3) and ABA signal pathways were significantly increased under salt stress. The expression of PnLRR-RLK27 significantly enhanced the resistance to salt stress and the sensitivity to ABA in the gametophyte growth stage. The function and mechanism of the Arabidopsis in Arabidopsis thaliana: the pROK2 vector of PnLRR RLK27 was constructed and the Arabidopsis flower was infected by Agrobacterium tumefaciens, and the pure lines of the over expressed PnLRR-RLK27 transgenic Arabidopsis were obtained. Under salt stress, the seed germination rate of the transgenic plant was 1.5 times higher than that of the wild type plant. The main root of the transgenic plant was about 1.3 times.ABA of the wild type plant. The germination rate of the transgenic plants was 2 times that of the wild type, and the main root was 10 times longer than that of the wild type. Under the H202 treatment, the main root of the transgenic plant was 1. longer than the wild type. 4 times and 1.6 times the lateral root number, indicating that overexpression of PnLRR-RLK27 increased the resistance to salt stress and oxidative stress in Arabidopsis and reduced sensitivity to ABA. Compared with wild Arabidopsis, overexpression of PnLRR-RLK27 under salt treatment reduced the content of H202 and MDA, increased the content of prolysine and increased the antioxidant enzyme superoxide dismutase. (superoxide dismutase, SOD), the activity of peroxidase (peroxidase, POD) and catalase (catalase, CAT), up regulation of the expression level of the antioxidant enzyme gene At4PX1, AtAPX2, AtCAT2 and AtZAT10. The expression of 1 and AtCOR47 was significantly higher in the transgenic plants than in the wild type. The above results showed that the function of the PnLRR-RLK27 gene was conservative, and the function was similar in the two pattern plants, which could significantly improve the plant resistance to salt stress. The mechanism of its action may be to improve the scavenging ability of plant to ROS and the signal of stress related hormone ABA signal. .1.2 PnLRR-RLK19 enhanced the subcellular location analysis of plant resistance to salt stress and drought stress. It was found that the distribution of PnLRR-RLK19 in the cell membrane was overexpressed in Arabidopsis thaliana, and did not affect the normal growth of plants. After salt treatment, the germination rate of transgenic plants was 25% higher than that of wild type plants, and the main transgenic plants were dominant. The root of the plant was 2 times longer than that of the wild type. Under the treatment of 0.5 MABA, the seed germination rate of transgenic plants was more than 11.5% higher than that of the wild type plants. After 7.5 u M ABA treatment, the main roots of the transgenic plants were 1.5 times.H202 and mannitol treatment, and the number of overexpressed plants was 1.5 times and 2.0-4.0 times more than that of the wild type plants. After salt treatment, the leaf yellow of transgenic plants was less than that of wild type Arabidopsis. After drought treatment, the water loss rate of transgenic plants was low. After salt stress treatment, the ABA signal pathway genes (AtAREB1, AtAREB2, AtABF3, AtMYB2, AtDREB2A, AtRD22 and AtRD29A) and salt stress response genes (AtP5CS1 and AtSOS3) were significantly up-regulated after salt stress treatment. After drought treatment, the ABA signaling pathway genes of transgenic plants (AtABF3, AtMYB2, AtDREB2A, AtRD22, AtRD29A, AtRD29 and AtKIN1) showed obvious up-regulated expression patterns. The results showed that PnLRR-RLK19 not only enhanced the resistance of plants to salt stress, but also enhanced plant resistance to drought stress and oxidative stress; its mechanism and salt were also enhanced. The stress signal pathway and the positive correlation of the ABA signal pathway.1.3 PnLRR-RLK44 enhanced the plant's resistance to salt stress and abscisic acid in the cell membrane. The overexpression vector of PnLRR-RLK4 was constructed, the Arabidopsis thaliana was infected with Agrobacterium tumefaciens, and the transgenic homozygous plants were overexpressed. On the 100 mMNaCl medium. The germination rate of overexpressed plants was 17.6% higher than that of wild type plants. The seed germination rate of transgenic plants was 20% higher than that of wild type, and the main root of transgenic plants was 20% higher than that of wild type, and the main root of the transgenic plant was 20% higher than that of wild type, and the main root of the transgenic plant was 20% higher than that of the wild type. The signal pathway of salt stress and the related threat in the ABA pathway were detected by Real-time qPCR technology. In response to the expression of response genes, Salt Stressed ion channel protein genes (AtHKT1 and AtSOS3) were overexpressed. Salt stress response genes (AtP5CS1 and AtADH1) and ABA Insensitive Gene ABI1 were all significantly up-regulated. At the same time, the key genes of ABA synthesis pathway (AtNCED3, AtABA1 and AtAAO3) and ABA early response genes were found. Both RD29A and AtDREB2A were downregulated. These results suggest that PnLRR-RLK44 improves salt tolerance by participating in salt stress signaling pathways and with negative regulatory factors of ABA signaling pathways. In summary, PnLRR-RLK27 increases plant resistance to salt stress and reduces sensitivity to ABA. The mechanism may be associated with increasing plant resistance. The scavenging ability of ROS and the salt stress of /ABA signal pathway related.PnLRR-RLK19 not only improve plant resistance to salt stress and ABA insensitivity, but also enhance plant resistance to drought stress and oxidative stress, salt stress, osmotic stress, and ABA signal pathway related gene expression up regulation of.PnLRR-RLK44 through participation in salt stress signal pathway and A The negative regulatory factor of the BA signal pathway improves the salt tolerance of plants, the ABA synthesis and response pathway is down, and the negative regulation of ABA is a subfamily of.PnLRR-RLK09/27/44. The function of the gene is similar, but it also has some differentiation, which has its own uniqueness in improving the resistance mechanism of plants,.2 a ubiquitin. The role of enzyme in the response of Antarctic moss to abiotic stresses E3 ubiquitin ligase (E3 ubiquitin ligase, E3) can identify target proteins in a special way and play an important role in plant growth and response to abiotic stress. However, the study of the role of ubiquitin ligase in lower plants has not yet been reported. A ubiquitin ligase gene was cloned from the moss of Antarctica. The cDNA, named PnE3.PnE3, contains 1 2097 BP open reading frames, encoding 698 amino acids, and the molecular weight is 76.2 kDa. The isoelectric point is found to be 6.79. amino acid sequence analysis. PnE3 contains 1 Ubox domains (288-351 of aminoacid) and 4 ARM repeat domains (amino acid 418-625). Low temperature, high salt, drought and ABA can all induce the up-regulated gene expression, indicating that PnE3 gene participates in the response of Antarctic moss to abiotic stress. The transient expression vector of PnE3:GFP is converted to Arabidopsis thaliana. It is found that the green fluorescence of PnE3:GFP is mainly distributed in the cytoplasm, indicating that PnE3 is located in the cytoplasm. The function and mechanism in the small erect moss Under the treatment of 125 mM NaCl, the gametophyte diameter of wild type plants was 1.4 times larger than that of transgenic plants. Under 5 u M ABA treatment, the gametophyte diameter of wild type plants was 1.6 times larger than that of transgenic plants,.ABA signal pathway gene PpAB13A, PpABI3B and PpABI3C were down down in the transgenic small erect moss. The cotyledon opening rate of wild type plants was 40% higher than that of overexpression line, and the root length of wild type plant was 2.4 times longer than that of overexpression line. The cotyledon opening rate of wild type plants was 25.6% higher than that of overexpression line, and the main root of wild type plants was 3.7 times longer than that of overexpression line. Under salt stress, the stress response gene AtABI3, AtABFA, AtDREB2A and AtRD29A were under salt stress. The expression level of transgenic Arabidopsis was significantly downregulated. In conclusion, PnE3 gene is conservative in function, and can enhance plant pairs in two kinds of plants.
【学位授予单位】：山东大学
【学位级别】：博士
【学位授予年份】：2017
【分类号】：Q943.2

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