食烟性夜蛾科昆虫对生物碱的适应及仿生型信号分子对烟草生物碱的抑制
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摘要
本研究用烟草为模式植物,建立了烟草中四种主要生物碱及尼古丁的代谢物—可替宁的分离分析的常规方法。以斜纹夜蛾、烟青虫和棉铃虫三种夜蛾科昆虫为模式昆虫,通过测定它们取食烟叶后的虫体、虫粪,比较分析三种夜蛾科昆虫对烟草生物碱的适应机制。昆虫取食或机械性损伤都能激活烟草植株的系统性防卫反应,诱导根部尼古丁运输至受伤及相邻部位,正是在烟草与昆虫互作研究的基础上,选取了一种与烟株创伤信号相颉抗的仿生型信号分子(BSM),研究其与品种、施肥量以及打顶时期等栽培因素的互作效应。优化BSM的应用,从而将烟草生物碱总量及生物碱组成控制在合适的范围,为烟草生物碱的调控提供一条新途径。
论文主要工作总结如下:
1.通过大量实验,探索出了一种简单、快速和可靠的反相高效液相色谱法测定烟草中降烟碱、新烟草碱、假木贼碱、尼古丁及其代谢物可替宁的含量。系统的优化了前处理过程中萃取溶剂和萃取方式,应用C18固相小柱进行样品的纯化。通过优化色谱条件最终确定了最佳的仪器测定条件,并从线性范围、精密度和回收率等方面对分析方法进行了验证。实验结果表明,方法具有较宽的线性范围和较高的回收率,五种物质的相对标准偏差在0.89-3.56%之间,整个实验过程加标回收率均大于90%,适合于烟草中生物碱的分析测定。
2.烟草是斜纹夜蛾、烟青虫和棉铃虫的共同寄主,利用反相高效液相色谱法测定虫体、虫粪和取食的烟叶中生物碱和可替宁的含量以确定昆虫摄取的生物碱是否被吸收、截留、代谢和排泄,从而确定三种夜蛾科昆虫对生物碱的适应机制。结果表明,只有少量生物碱被截留和解毒成可替宁,大部分生物碱是以原来的形式被排出体外。而且,烟青虫和棉铃虫具有比斜纹夜蛾更强的尼古丁代谢能力和生物碱排泄能力。虫粪放置后,可替宁的含量增加,而生物碱含量则降低,表明虫粪中的生物碱可以在外界环境如空气或微生物条件下自然降解。三种夜蛾科昆虫之所以能适应烟草作为寄主植物主要是因为他们拥有强有效的排泄机制。
3.用源于取食烟草的寡食性昆虫口腔分泌物的仿生型信号分子(BSM)在不同条件下、对不同类型烟草在打顶后进行涂抹处理。用HPLC检测BSM处理株和对照株烟草主要生物碱的含量,分析探讨了BSM在不同条件下对烟草生物碱的抑制作用。结果表明:BSM物质对不同类型烟草打顶后叶片上生物碱的积聚均有不同程度的抑制作用。BSM对烤烟9717中降烟碱的影响最大,中部和上部叶分别降低55.78 %和63.40 %,上部叶各生物碱降幅均大于中部叶,其中尼古丁和总生物碱分别降低25.49 %和26.28 %。BSM浓度对白肋烟中生物碱的抑制效应也有影响,0.1 mol/L的BSM对尼古丁的影响最大,中、上部叶的降低率分别达22.99 %和42.45 %,总生物碱则分别降低22.21 %和37.73 %;0.2 mol/L的BSM可使降烟碱的量显著降低,中部和上部叶分别降低62.94 %和58.53 %。施氮量对BSM的抑制作用也有影响,在高的施氮量(9 kgN/667 m2)下抑制作用更大,可使K326上部叶总生物碱降低34.76 %。
4.研究了烤烟云烟85在7kg N/667m2和9kg N/667m2施氮量时,在现蕾、初花、中花和盛花四个不同花期打顶后施用BSM对烟株中各生物碱含量的影响。结果表明,在两种施氮量时,BSM对不同花期处理的烟株中生物碱的积累均有抑制作用。在现蕾期处理可有效抑制尼古丁积累,在中花期处理可显著降低三种微量生物碱积累。就两种施氮量而言,在9kg N/667m2时各花期处理后的烟株中生物碱含量与对照相比降幅较大。
This paper seeked to explore a simple routine analytical method to separate and determine four major alkaloids in tobacco and cotinine of metabolism of nicotine, and apply this methodology to analyze alkaloids and cotinine of larvae, frass, and tobacco, in order to determine the adaptation mechanism of noctuid caterpillars to tobacco alkaloids. Herbivore feeding or mechanical wounding can activate some chemical defense system of tobacco, which induce a rapid increase of nicotine in roots and transport to damaged and adjacent parts. On the basis of the interaction between tobacco and insects, a bionics signal molecule which has a gainst responses to would signal of tobacco was selected to treated tobacco. The research on the interaction effects between bionics signal molecule and the cultivation factors of varieties, fertilizer, and topping time were studied. The contents of tobacco alkaloids and composition were controlled in a appropriate range by optimizing the application conditions of BSM. To provide a new way in the control of alkaloids content in tobacco leaves.
Four main aspects were included in the dissertation:
1. A novel, simple, quick and reliable method for analysis of nornicotine, anatabine, anabasine, nicotine and cotinine has been developed utilizing reverse high performance liquid chromatography. A series of optimization were done to find the most effective extraction solvent and technology for tobacco sample, and application of solid-phase C18 column for purification of samples. The best determination conditions were chosen after a series of optimization, and the method was validated by linearity, percision, repeatability and recovery. RSD value of five reference materials were from 0.89 to 3.56%, and the overall recovery was above 90%, all these datas showed that this improved method was feasible for tobacco analysis system.
2. Tobacco is a common host plant of Helicoverpa armigera (Hübner), Helicoverpa assulta (Guenée) and Spodoptera litura (Fabricius) (Lepidoptera, Noctuidae). Larvae, frass and tobacco were analyzed by HPLC to determine if alkaloids were assimilated, sequestered, metabolized and excreted. In order to confirm the adaptation mechanism of this three noctuid caterpillars to tobacco alkaloids. The results of this investigation strongly suggest that only a small amount of alkaloids are sequestered and detoxified into cotinine by three noctuid insects, and alkaloids are excreted unmodified. Furthermore, H.assulta and H.armigera have stronger ability of nicotine metabolism and alkaloids excretion capacity than S.litura. Cotinine concentration in storaged frass of S.litura were increased with the growth of storage time, and alkaloids content were declined in this conditions. It is showed that alkaloids contained in the frass naturally degrades to cotinine in the presence of environmental conditions such as air , or microorganism. Three noctuid insects which have become adapted to tobacco as a host plant mainly because of an efficient excretory mechanism.
3. The application of bionics signal molecule (BSM) from tobacco feeding oligophagous insect oral secretion to diverse types of tobacco after topping in different conditions. The major alkaloids concentrations in tobacco leaves treated with BSM and untreated control were analyzed by HPLC, and the impact of BSM on the alkaloids under different conditions were discussed. Results indicated that BSM could inhibit the alkaloids accumulation after topping in various tobacco leaves at different levels. Applied BSM on 9717 could markedly reduced the level of nornicotine in middle and upper leaves by 55.78% and 63.40% respectively, while alkaloids in upper leaves decrease more in comparison with middle leaves, there were 25.49% and 26.28% decline of nicotine and total alkaloids in upper leaves. The concentrations of BSM also have inhibitory effects on alkaloids accumulation in burley, with the biggest effect on nicotine by 0.1 mol/L and nornicotine by 0.2 mol/L. Applied 0.1 mol/L BSM to burley displayed a decrease of nicotine by 22.99% and 42.45%, total alkaloids by 22.21% and 37.73% in middle and upper leaves respectively ,While treated with 0.2 mol/L BSM showed a decrease of nornicotine by 62.94% and 58.53 % in middle and upper leaves respectively. Furthermore, there are some influence of nitrogen content on the regulating effect of BSM in K326 . The total alkaloids concentrations of upper leaves were lowest with 9 kg/667 m2 N applied , an decrease of 34.76 % compared with the control.
4. The effects of the topping time in the budding, early-flowering, medium- flowering and full-blooming stage treated with BSM at nitrogen rate of 7kg N/667m2 and 9kg N/667m2 on the content of alkaloids in flue-cured tobacco yunyan 85 were studied, which indicated that both the early and late topping treated by BSM could inhibit the alkaloids accumulation in tobacco leaves. Tobacco treated by BSM aftertopping at budding stage were favorable for reducing the nicotine accumulation and the topping at medium-flowering stage could dramatically decrease the three minor alkaloids accumulation in tobacco leaves. The content of alkaloids in different topping time treated with BSM at nitrogenous fertilizer of 9kg N/667m2 has greater decline compared to 7kg N/667m2.
论文主要工作总结如下:
1.通过大量实验,探索出了一种简单、快速和可靠的反相高效液相色谱法测定烟草中降烟碱、新烟草碱、假木贼碱、尼古丁及其代谢物可替宁的含量。系统的优化了前处理过程中萃取溶剂和萃取方式,应用C18固相小柱进行样品的纯化。通过优化色谱条件最终确定了最佳的仪器测定条件,并从线性范围、精密度和回收率等方面对分析方法进行了验证。实验结果表明,方法具有较宽的线性范围和较高的回收率,五种物质的相对标准偏差在0.89-3.56%之间,整个实验过程加标回收率均大于90%,适合于烟草中生物碱的分析测定。
2.烟草是斜纹夜蛾、烟青虫和棉铃虫的共同寄主,利用反相高效液相色谱法测定虫体、虫粪和取食的烟叶中生物碱和可替宁的含量以确定昆虫摄取的生物碱是否被吸收、截留、代谢和排泄,从而确定三种夜蛾科昆虫对生物碱的适应机制。结果表明,只有少量生物碱被截留和解毒成可替宁,大部分生物碱是以原来的形式被排出体外。而且,烟青虫和棉铃虫具有比斜纹夜蛾更强的尼古丁代谢能力和生物碱排泄能力。虫粪放置后,可替宁的含量增加,而生物碱含量则降低,表明虫粪中的生物碱可以在外界环境如空气或微生物条件下自然降解。三种夜蛾科昆虫之所以能适应烟草作为寄主植物主要是因为他们拥有强有效的排泄机制。
3.用源于取食烟草的寡食性昆虫口腔分泌物的仿生型信号分子(BSM)在不同条件下、对不同类型烟草在打顶后进行涂抹处理。用HPLC检测BSM处理株和对照株烟草主要生物碱的含量,分析探讨了BSM在不同条件下对烟草生物碱的抑制作用。结果表明:BSM物质对不同类型烟草打顶后叶片上生物碱的积聚均有不同程度的抑制作用。BSM对烤烟9717中降烟碱的影响最大,中部和上部叶分别降低55.78 %和63.40 %,上部叶各生物碱降幅均大于中部叶,其中尼古丁和总生物碱分别降低25.49 %和26.28 %。BSM浓度对白肋烟中生物碱的抑制效应也有影响,0.1 mol/L的BSM对尼古丁的影响最大,中、上部叶的降低率分别达22.99 %和42.45 %,总生物碱则分别降低22.21 %和37.73 %;0.2 mol/L的BSM可使降烟碱的量显著降低,中部和上部叶分别降低62.94 %和58.53 %。施氮量对BSM的抑制作用也有影响,在高的施氮量(9 kgN/667 m2)下抑制作用更大,可使K326上部叶总生物碱降低34.76 %。
4.研究了烤烟云烟85在7kg N/667m2和9kg N/667m2施氮量时,在现蕾、初花、中花和盛花四个不同花期打顶后施用BSM对烟株中各生物碱含量的影响。结果表明,在两种施氮量时,BSM对不同花期处理的烟株中生物碱的积累均有抑制作用。在现蕾期处理可有效抑制尼古丁积累,在中花期处理可显著降低三种微量生物碱积累。就两种施氮量而言,在9kg N/667m2时各花期处理后的烟株中生物碱含量与对照相比降幅较大。
This paper seeked to explore a simple routine analytical method to separate and determine four major alkaloids in tobacco and cotinine of metabolism of nicotine, and apply this methodology to analyze alkaloids and cotinine of larvae, frass, and tobacco, in order to determine the adaptation mechanism of noctuid caterpillars to tobacco alkaloids. Herbivore feeding or mechanical wounding can activate some chemical defense system of tobacco, which induce a rapid increase of nicotine in roots and transport to damaged and adjacent parts. On the basis of the interaction between tobacco and insects, a bionics signal molecule which has a gainst responses to would signal of tobacco was selected to treated tobacco. The research on the interaction effects between bionics signal molecule and the cultivation factors of varieties, fertilizer, and topping time were studied. The contents of tobacco alkaloids and composition were controlled in a appropriate range by optimizing the application conditions of BSM. To provide a new way in the control of alkaloids content in tobacco leaves.
Four main aspects were included in the dissertation:
1. A novel, simple, quick and reliable method for analysis of nornicotine, anatabine, anabasine, nicotine and cotinine has been developed utilizing reverse high performance liquid chromatography. A series of optimization were done to find the most effective extraction solvent and technology for tobacco sample, and application of solid-phase C18 column for purification of samples. The best determination conditions were chosen after a series of optimization, and the method was validated by linearity, percision, repeatability and recovery. RSD value of five reference materials were from 0.89 to 3.56%, and the overall recovery was above 90%, all these datas showed that this improved method was feasible for tobacco analysis system.
2. Tobacco is a common host plant of Helicoverpa armigera (Hübner), Helicoverpa assulta (Guenée) and Spodoptera litura (Fabricius) (Lepidoptera, Noctuidae). Larvae, frass and tobacco were analyzed by HPLC to determine if alkaloids were assimilated, sequestered, metabolized and excreted. In order to confirm the adaptation mechanism of this three noctuid caterpillars to tobacco alkaloids. The results of this investigation strongly suggest that only a small amount of alkaloids are sequestered and detoxified into cotinine by three noctuid insects, and alkaloids are excreted unmodified. Furthermore, H.assulta and H.armigera have stronger ability of nicotine metabolism and alkaloids excretion capacity than S.litura. Cotinine concentration in storaged frass of S.litura were increased with the growth of storage time, and alkaloids content were declined in this conditions. It is showed that alkaloids contained in the frass naturally degrades to cotinine in the presence of environmental conditions such as air , or microorganism. Three noctuid insects which have become adapted to tobacco as a host plant mainly because of an efficient excretory mechanism.
3. The application of bionics signal molecule (BSM) from tobacco feeding oligophagous insect oral secretion to diverse types of tobacco after topping in different conditions. The major alkaloids concentrations in tobacco leaves treated with BSM and untreated control were analyzed by HPLC, and the impact of BSM on the alkaloids under different conditions were discussed. Results indicated that BSM could inhibit the alkaloids accumulation after topping in various tobacco leaves at different levels. Applied BSM on 9717 could markedly reduced the level of nornicotine in middle and upper leaves by 55.78% and 63.40% respectively, while alkaloids in upper leaves decrease more in comparison with middle leaves, there were 25.49% and 26.28% decline of nicotine and total alkaloids in upper leaves. The concentrations of BSM also have inhibitory effects on alkaloids accumulation in burley, with the biggest effect on nicotine by 0.1 mol/L and nornicotine by 0.2 mol/L. Applied 0.1 mol/L BSM to burley displayed a decrease of nicotine by 22.99% and 42.45%, total alkaloids by 22.21% and 37.73% in middle and upper leaves respectively ,While treated with 0.2 mol/L BSM showed a decrease of nornicotine by 62.94% and 58.53 % in middle and upper leaves respectively. Furthermore, there are some influence of nitrogen content on the regulating effect of BSM in K326 . The total alkaloids concentrations of upper leaves were lowest with 9 kg/667 m2 N applied , an decrease of 34.76 % compared with the control.
4. The effects of the topping time in the budding, early-flowering, medium- flowering and full-blooming stage treated with BSM at nitrogen rate of 7kg N/667m2 and 9kg N/667m2 on the content of alkaloids in flue-cured tobacco yunyan 85 were studied, which indicated that both the early and late topping treated by BSM could inhibit the alkaloids accumulation in tobacco leaves. Tobacco treated by BSM aftertopping at budding stage were favorable for reducing the nicotine accumulation and the topping at medium-flowering stage could dramatically decrease the three minor alkaloids accumulation in tobacco leaves. The content of alkaloids in different topping time treated with BSM at nitrogenous fertilizer of 9kg N/667m2 has greater decline compared to 7kg N/667m2.
引文
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