三种植物凝集素基因在转基因小麦中的表达及其抗蚜效果分析
摘要
小麦生产在国家粮食安全中占据举足轻重的地位,近年来,随着麦田水肥条件的改善和农业生态环境的变化,麦蚜危害日趋加重。常规育种在抗蚜小麦种质资源筛选与鉴定方面作了大量工作,但进展缓慢。利用转基因技术可弥补传统小麦育种的不足,突破可利用基因库的限制。植物凝集素对具有刺吸式口器的麦蚜具有较好的抗性,可以有效抑制其生长和繁殖率,因此通过转基因技术将其导入普通小麦品种,培育小麦抗蚜新种质,从而提高我国小麦的抗蚜水平。
本研究根据小麦基因密码子的偏好性和天然凝集素的氨基酸信息,人工合成了雪花莲凝集素基因(synthesized Galanthus navalis agglutinin, sGNA)与中国水仙凝集素基因(synthesized Narcissus tazetta lectin,sNTL),并通过RT-PCR获得了半夏凝集素基因(Pinellia ternate agglutinin, pta),分别构建了由番茄核酮糖-1、5-二磷酸羧化酶小亚基基因(rbcs)启动子驱动的3个表达载体,通过基因枪法转化普通小麦品种,以研究其在小麦中的表达和抗蚜效果。主要研究结果如下：
(1)通过转化小麦幼胚愈伤组织,经分化和再生,分别获得42株、65株、54株T0代阳性植株。对后代转基因植株进行PCR、RT-PCR、Southern blot、Northern blot、以及转基因株系的叶片蛋白提取液的凝血活性的检测,确认sGNA与sNTL基因成功整合到小麦基因组中并且能够稳定表达,分别筛选出12个与11个T4代纯合株系。
(2)通过分析株系zy2-18外源基因插入位点的边界序列,初步判断转sGNA基因株系2-18的外源片段是通过同源重组和随机插入的方式整合到小麦A染色体组。
(3)对转sGNA与sNTL基因纯合株系的抗蚜性分别进行室内离体叶片鉴定以及两年的田间鉴定,结果表明5个(zy2-11、zy2-18、jd8-10、yn5-2、yn5-20)转sGNA基因小麦纯合株系与7个(fy4-4、fy4-10、af7-8、af7-10、zy9-1、zy9-4、zy9-8)转sNTL基因纯合株系的蚜虫致死率、增长率以及单分蘖蚜虫发生量与相应对照品种差异显著或极显著。且转sGNA株系yn5-20抗虫水平提高幅度最高,可达75.6%；在转sNTL基因株系中af7-8的抗虫水平比对照AF9高出80.3%。另外经室内离体叶片鉴定,6株转pta基因小麦表现出了较高的抗蚜水平。
(4)转sGNA基因株系及相应对照的单分蘖蚜虫相对发生量与叶片蛋白提取液的OD值之间呈显著正相关(r=0.812*)关系,转sNTL基因株系及相应对照的单分蘖蚜虫相对发生量与叶片蛋白提取液的OD值之间呈极显著正相关(r=0.0.932*)关系。因此,转基因株系抗虫水平的提高主要原因在于sGNA与sNTL蛋白的表达。
上述结果表明,转sGNA、sNTL和pta基因小麦都具有较好的抗蚜虫效果,因此,可以将它们作为培育抗蚜小麦的优良外源基因,并聚合多个基因的以进一步提高转基因小麦的抗蚜性。
Wheat has a special status in stabilizing China food supply. The product loss of wheat due to insects is very high in recent years. For a long time the control of these pests has depended chiefly on large amounts of insecticides sprays which cause considerable environmental pollution and represent a health hazard to farmers as well as significantly increasing the costs of wheat production. Conventional plant breeding programs have been undertaken and considerable time and effort has been made in searching for insect-resistant germplasm resources, but it has moved slowly. Genetically modified (gm) can make up the deficiency of the conventional wheat breeding and breakthrough the limitation of available gene pool. Protease inhibitors, Bt endotoxins and plant lectins have been proven to be very successful in control insects. A more complete transformation system of particle bombardment method has been established, with young embryo and young ear as the receptor, which made it is possible to fostering high-quality and aphid-resistant wheat by genetic engineering.
Plant lectins had been shown toxic to sucking pests resulting in reduced survival and delayed development. It's significantly for developing aphid-resistant wheat germplasm resources by introducing these genes into common wheat.
In this study, sGNA (synthesized Galanthus nivalis agglutinin) and sNTL (synthesized Narcissus tazetta lectin) genes encoding mannose binding lectins were synthesized according to the codons preferable principle of wheat genes and transgenic sGNA and sNTL common wheat lines, driven by constitutive and phloem-specific promoter rbcs, were generated by particle bombardment transformation. The objectives of this study were:(1) to confirm that the two synthetic genes sGNA and sNTL can be effectively expressed in transgenic wheat respectively;(2) to examine the aphid resistance performance of sGNA transgenic lines and sNTL transgenic lines indoors as well as field conditions and evaluate the functions of sGNA and sNTL against wheat aphid; and (3) to produce transgenic lines that may be useful for developing aphid-resistant wheat germplasm resources. Main research results are as follows:
(1)42To sGNA transgenic plants,56To sNTL transgenic plants and54To pta transgenic plants were obtained by particle bombardment transformation with embryogenic calli as receptor. Twelve homozygous T4sGNA transgenic lines and eleven homozygous T4sNTL transgenic lines were identified by PCR amplification. The three genes were successfully integrated into the wheat genome and expressed stable according to the tests of PCR, RT-PCR、Southern blot. Northern blot and clotting test.
(2) The transgene insert site in single gene patental line zy2-18containing sGNA were analyzed preliminary. The flanking sequences of sGNA were cloned by genomewalking, and located on chromosome A by the homologous recombination and random insertion.
(3) After the insect bioassays in the lab and field-test, five sGNA transgenic lines (zy2-11、zy2-18、 jd8-10、yn5-2、yn5-20) and seven sNTL transgenic lines (fy4-4、fy4-10、af7-8、af7-10、zy9-1、 zy9-4、zy9-8) were found highly resistant to aphids compared to the corresponding control plants. Significant differences were also observed for mortality and growth rate of aphids between the transgenic lines and the corresponding controls in lab condition on detached leaf tissues. The aphid amount per tiller of af7-8sNTL transgenic line was reduced by80.3%maximum compared to the control Af9. Six pta transgenic plants showed a higher level of resistance genes in wheat□ccordingto the identification of detached leaf segment.
(4) The sGNA transgenic lines and the controls showed a significant positive correlation (r=0.812*) between the relative aphid amount per tiller and OD values of leaf tissues extracts. The sNTL transgenic lines, of which it showed a significant positive correlation (r=0.932**) between aphid amount per tiller and OD values of leaf tissues extracts. So the result of clotting activity test indicated that the increased resistance to aphids was significantly correlated with the expression of sGNA and sNTL lectins in the transgenic lines.
Therefore pta, sGNA and sNTL could be served as alternative choice to produce aphid-resistance wheat through genetically modified approach. It may also be worth commenting on the strategies for pyramiding the lectin genes.
本研究根据小麦基因密码子的偏好性和天然凝集素的氨基酸信息,人工合成了雪花莲凝集素基因(synthesized Galanthus navalis agglutinin, sGNA)与中国水仙凝集素基因(synthesized Narcissus tazetta lectin,sNTL),并通过RT-PCR获得了半夏凝集素基因(Pinellia ternate agglutinin, pta),分别构建了由番茄核酮糖-1、5-二磷酸羧化酶小亚基基因(rbcs)启动子驱动的3个表达载体,通过基因枪法转化普通小麦品种,以研究其在小麦中的表达和抗蚜效果。主要研究结果如下：
(1)通过转化小麦幼胚愈伤组织,经分化和再生,分别获得42株、65株、54株T0代阳性植株。对后代转基因植株进行PCR、RT-PCR、Southern blot、Northern blot、以及转基因株系的叶片蛋白提取液的凝血活性的检测,确认sGNA与sNTL基因成功整合到小麦基因组中并且能够稳定表达,分别筛选出12个与11个T4代纯合株系。
(2)通过分析株系zy2-18外源基因插入位点的边界序列,初步判断转sGNA基因株系2-18的外源片段是通过同源重组和随机插入的方式整合到小麦A染色体组。
(3)对转sGNA与sNTL基因纯合株系的抗蚜性分别进行室内离体叶片鉴定以及两年的田间鉴定,结果表明5个(zy2-11、zy2-18、jd8-10、yn5-2、yn5-20)转sGNA基因小麦纯合株系与7个(fy4-4、fy4-10、af7-8、af7-10、zy9-1、zy9-4、zy9-8)转sNTL基因纯合株系的蚜虫致死率、增长率以及单分蘖蚜虫发生量与相应对照品种差异显著或极显著。且转sGNA株系yn5-20抗虫水平提高幅度最高,可达75.6%；在转sNTL基因株系中af7-8的抗虫水平比对照AF9高出80.3%。另外经室内离体叶片鉴定,6株转pta基因小麦表现出了较高的抗蚜水平。
(4)转sGNA基因株系及相应对照的单分蘖蚜虫相对发生量与叶片蛋白提取液的OD值之间呈显著正相关(r=0.812*)关系,转sNTL基因株系及相应对照的单分蘖蚜虫相对发生量与叶片蛋白提取液的OD值之间呈极显著正相关(r=0.0.932*)关系。因此,转基因株系抗虫水平的提高主要原因在于sGNA与sNTL蛋白的表达。
上述结果表明,转sGNA、sNTL和pta基因小麦都具有较好的抗蚜虫效果,因此,可以将它们作为培育抗蚜小麦的优良外源基因,并聚合多个基因的以进一步提高转基因小麦的抗蚜性。
Wheat has a special status in stabilizing China food supply. The product loss of wheat due to insects is very high in recent years. For a long time the control of these pests has depended chiefly on large amounts of insecticides sprays which cause considerable environmental pollution and represent a health hazard to farmers as well as significantly increasing the costs of wheat production. Conventional plant breeding programs have been undertaken and considerable time and effort has been made in searching for insect-resistant germplasm resources, but it has moved slowly. Genetically modified (gm) can make up the deficiency of the conventional wheat breeding and breakthrough the limitation of available gene pool. Protease inhibitors, Bt endotoxins and plant lectins have been proven to be very successful in control insects. A more complete transformation system of particle bombardment method has been established, with young embryo and young ear as the receptor, which made it is possible to fostering high-quality and aphid-resistant wheat by genetic engineering.
Plant lectins had been shown toxic to sucking pests resulting in reduced survival and delayed development. It's significantly for developing aphid-resistant wheat germplasm resources by introducing these genes into common wheat.
In this study, sGNA (synthesized Galanthus nivalis agglutinin) and sNTL (synthesized Narcissus tazetta lectin) genes encoding mannose binding lectins were synthesized according to the codons preferable principle of wheat genes and transgenic sGNA and sNTL common wheat lines, driven by constitutive and phloem-specific promoter rbcs, were generated by particle bombardment transformation. The objectives of this study were:(1) to confirm that the two synthetic genes sGNA and sNTL can be effectively expressed in transgenic wheat respectively;(2) to examine the aphid resistance performance of sGNA transgenic lines and sNTL transgenic lines indoors as well as field conditions and evaluate the functions of sGNA and sNTL against wheat aphid; and (3) to produce transgenic lines that may be useful for developing aphid-resistant wheat germplasm resources. Main research results are as follows:
(1)42To sGNA transgenic plants,56To sNTL transgenic plants and54To pta transgenic plants were obtained by particle bombardment transformation with embryogenic calli as receptor. Twelve homozygous T4sGNA transgenic lines and eleven homozygous T4sNTL transgenic lines were identified by PCR amplification. The three genes were successfully integrated into the wheat genome and expressed stable according to the tests of PCR, RT-PCR、Southern blot. Northern blot and clotting test.
(2) The transgene insert site in single gene patental line zy2-18containing sGNA were analyzed preliminary. The flanking sequences of sGNA were cloned by genomewalking, and located on chromosome A by the homologous recombination and random insertion.
(3) After the insect bioassays in the lab and field-test, five sGNA transgenic lines (zy2-11、zy2-18、 jd8-10、yn5-2、yn5-20) and seven sNTL transgenic lines (fy4-4、fy4-10、af7-8、af7-10、zy9-1、 zy9-4、zy9-8) were found highly resistant to aphids compared to the corresponding control plants. Significant differences were also observed for mortality and growth rate of aphids between the transgenic lines and the corresponding controls in lab condition on detached leaf tissues. The aphid amount per tiller of af7-8sNTL transgenic line was reduced by80.3%maximum compared to the control Af9. Six pta transgenic plants showed a higher level of resistance genes in wheat□ccordingto the identification of detached leaf segment.
(4) The sGNA transgenic lines and the controls showed a significant positive correlation (r=0.812*) between the relative aphid amount per tiller and OD values of leaf tissues extracts. The sNTL transgenic lines, of which it showed a significant positive correlation (r=0.932**) between aphid amount per tiller and OD values of leaf tissues extracts. So the result of clotting activity test indicated that the increased resistance to aphids was significantly correlated with the expression of sGNA and sNTL lectins in the transgenic lines.
Therefore pta, sGNA and sNTL could be served as alternative choice to produce aphid-resistance wheat through genetically modified approach. It may also be worth commenting on the strategies for pyramiding the lectin genes.
引文
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