取食转基因棉的棉蚜对中华草蛉的影响及Bt毒蛋白在三级营养链中的传递
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摘要
本论文以生产上大规模商业化种植的转基因棉(国抗12,新棉33B和SGK321)及其对应常规棉(泗棉3号,新棉33,石远321)为材料,测定了各转基因棉不同生育期和不同部位的Bt毒蛋白含量；以取食各品种棉花的棉蚜(Aphis gossypii)饲喂中华草蛉(Chrysoperla sinica),研究不同品种转基因棉对中华草蛉生长发育与繁殖的间接影响以及中华草蛉对各品种棉田棉蚜的捕食功能反应及搜寻效应；依托生态系统的重要构成部分——食物链,研究Bt毒蛋白在棉花-棉蚜-蜜露-针毛收获蚁(Messor aciculatus)中的转移,以及在棉花-棉蚜-中华草蛉中的传递。主要研究结果如下：
1.转基因抗虫棉Bt毒蛋白表达量的时空变化研究结果表明：Bt基因在所有转基因棉花器官中均有表达,不同生育期功能叶中的Bt毒蛋白含量差异显著,变化趋势为先高后低再略有回升,即苗期>蕾期>花期>吐絮期>铃期；蕾、花、铃各部位中的Bt毒蛋白含量的时间变化也有相同趋势。Bt毒蛋白在各时期的含量均表现为功能叶中最高,大于花、蕾等繁殖器官。不同叶位对Bt毒蛋白含量有直接影响。不同花器官中Bt毒蛋白含量有明显差异,表现为花瓣>花蕊>子房>苞片,其中苞片中Bt毒蛋白含量远远低于花瓣等其它花器官。GK12苞片中Bt毒蛋白含量最低,仅为147.9ng/g;最高Bt毒蛋白含量在SGK321功能叶中被检测到,达1630.1ng/g,为GK12苞片中含量的11.0倍。转基因棉叶中杀虫蛋白在土壤环境中的降解趋势表现为初期降解非常迅速,之后趋于平缓。试验开始的第一个月里(2010年9月～2010年10月),GK12、33B、SGK321三个品种中的杀虫蛋白分别降解了76.78%,82.03%和84.26%,至次年4月份,几乎检测不到Bt蛋白,到5月份已完全检测不到Bt蛋白存在。
2.取食不同品种棉花的蚜虫对中华草蛉生长发育及繁殖的影响结果指出：中华草蛉捕食不同品种棉花上的棉蚜后,其死亡率、羽化率以及茧的发育历期、茧重和成虫性比在棉花品种间大多无显著差异；取食SGK321棉田棉蚜的草蛉幼虫1代死亡率为8.89%,还极显著低于对照处理(40.0%),2代死亡率为16.09%,也显著低于对照处理(75.9%)；取食SGK321棉田棉蚜的草蛉幼虫1、2代羽化率较高,分别为对照品种的1.18倍和1.17倍。各处理间草蛉成虫的产卵前期无显著差异；产卵期表现为取食转基因棉田棉蚜的草蛉都显著长于对照处理,其中取食SGK321棉田棉蚜的草蛉1代产卵历期达33.01天,显著高于对照；用取食转基因棉的棉蚜饲喂草蛉,后者的产卵量高于对应处理。其中,取食转基因棉田棉蚜的草蛉1、2代单雌产卵量部分显著高于对照,且取食SGK321棉田棉蚜的草蛉幼虫1代单雌产卵量高达882粒,显著高于对照处理。各处理间草蛉卵的孵化率无显著差异。一系列研究结果均表明,转Bt基因抗虫棉对中华草蛉的生长发育和繁殖无不良影响。取食转基因棉,尤其是SGK321品种上的棉蚜对中华草蛉的生长发育与繁殖似有一定的促进作用。
3.中华草蛉对不同品种棉田棉蚜的功能反应与搜寻效应结果发现,草蛉幼虫取食蚜虫的数量随猎物密度的增加而增大,虽然草蛉对转基因棉田蚜虫的捕食量大多高于常规棉田,但差异不显著。草蛉幼虫各虫龄对棉蚜的捕食功能反应均符合Holling Ⅱ型。功能反应方程指出,草蛉幼虫的最大日捕食量常规棉略低于转基因棉,捕食转基因棉田棉蚜的处理时间短于捕食常规棉田的棉蚜。草蛉2龄、3龄幼虫对SGK321棉田棉蚜的日平均捕食量高于对照29%和41%。研究指出,草蛉各龄幼虫对常规棉田棉蚜的捕食作用率要低于转基因棉处理。草蛉各龄幼虫捕食转基因棉田棉蚜的种内干扰强度高于对照常规棉处理。分摊竞争强度研究结果表明：中华草蛉对转基因棉田棉蚜的分摊竞争强度大于常规棉田,1龄、2龄、3龄中华草蛉对转基因棉田棉蚜的最高分摊竞争强度分别高于对照品种棉田棉蚜的29.6%,27.4%和69.6%。由此可见,转基因棉对中华草蛉的捕食作用无不良影响。
4.Bt毒蛋白在棉花-棉蚜-蜜露-针毛收获蚁中的转移及在棉花-棉蚜-中华草蛉食物链中的传递结果发现：转基因棉叶中存在高浓度的Bt毒蛋白,三个转基因棉品种苗期功能叶的Bt毒蛋含量均达1000ng/g以上,而韧皮部汁液中的Bt毒蛋白含量非常低,苗期、花铃期分别仅为棉叶含量的0.36%和0.37%。取食转基因棉花的棉蚜体内也检测到了Bt毒蛋白,但含量甚微。连续、多代取食转基因棉的棉蚜,其体内Bt毒蛋白含量较高,是取食转基因棉一代棉蚜的1.16倍。表明随着取食代数的增加,毒蛋白在棉蚜体内似有一定的累积。取食各转基因棉的棉蚜分泌的蜜露量差异显著,常规棉田棉蚜泌蜜量达4.1g/d,是SGK321转基因棉棉蚜泌蜜量的2.28倍。取食转基因棉的棉蚜分泌的蜜露中检测到了Bt毒蛋白,但处理间无显著差异。围绕棉蚜的针毛收获蚁体内检测到微量的Bt毒蛋白。1代、2代中华草蛉幼虫捕食不同品种转基因棉田的棉蚜后,其体内均检测不出Bt毒蛋白,表明Bt毒蛋白不经过棉蚜传至其天敌中华草蛉体内。
Transgenic Bt cotton varieties have been commercialized in a large scale. In this study, transgenic cotton varieties (GK12, Xinmian33B, SGK321) and their corresponding conventional cotton controls (Simian3, Xinmian33and Shiyuan321) were used to determine Bt toxin content at different developmental stages and different parts of transgenic cotton. The effects of Aphis gossypii feeding in transgenic cotton on the growth, development, the reproduction, the predation functional responses and search effects of Chrysoperla sinica were evaluated. Transfer of Bt toxin protein among transgenic cotton-A. gossypii-honeydew and nature enemies was studied. The main results were as follows:
1. Temporal and spatial variation of the Bt toxin protein content in transgenic cotton showed that Bt gene was expressed in all organs, and there were significant differences in different growth stages of functional leaves. The variation trend was that the Bt expression was high and then decreased, and rose up a little at last, and seedling stage> bud stage> flowering> boll opening period>boll stage; the temporal variation of the Bt toxin protein content of bud, flower, bell also had similar trends. The spatial dynamics of Bt toxin proteins in the same period were consistent. The Bt toxin protein content of functional leaves was highest, and it was more than flowers, buds and other reproductive organs. Different leaf positions had a direct impact on the protein content of the Bt toxin. Bt toxin protein were significantly different in the different floral organs, that is, petal> flower> ovary> bract, and the Bt toxin protein in the bracts was far below the petals and other floral organs. Bt toxin protein in GK12bracts was the lowest, only147.9ng/g; highest Bt toxin protein was detected in SGK321functional leaves,1630.1ng/g,11.0times as the minimum content. In natural environment, the insecticidal protein in Bt cotton degraded much rapidly in the initial period and then slowered, it reached76.78%,82.03%and84.26%in first month (2010.9～2010.10) respectively in GK12, Xinmian33B, SGK321, and undetectable in late May.
2. Impact on the growth development and the reproduction of C. sinica feeding on A. gossypii in different varieties of cotton indicated that after preying on the A. gossypii on different varieties of cotton, the mortality rate, the emergence rate and cocoon developmental duration, cocoon weight and adult sex ratio of C. sinica in cotton varieties were almost no significant differences; only the mortality rate of the first generation of C. sinica larvae feeding on A. gossypii in SGK321was only8.89%, extremely lower than the control (40.0%). The mortality rate of the second generation was16.09%, significantly lower than the control (75.9%); and the emergence rate of the first and second generations of the C. sinica larvae feeding on the A. gossypii of SGK321was high,1.18and1.17times as the control. There was no significant difference among the treatments of C. sinica adult in preoviposition; The spawning period for C. sinica feeding on the A. gossypii of transgenic cotton were longer than the control; the oviposition duration of the first generation of C. sinica feeding on the A. gossypii in SGK321was up to33.01d, significantly longer than the control. The fecundity of C. sinica feeding on the A. gossypii in transgenic cotton was higher than the control. The fecundity of the first and second generation of the C. sinica feeding on the A. gossypii in33B and SGK321were significantly higher than control, and the fecundity of the first generation of C. sinica feeding on the A. gossypii in SGK321was up to882, which was extremely significantly higher than the control. There was no significant difference in egg hatchability between treatments. A series of studies showed that transgenic cotton had no adverse effects on the growth and reproduction of C. sinica. A. gossypii in transgenic cotton especially SGK321seems to have certain positive role in promoting the growth and development and reproduction of C. sinica.
3. Predation functional response and searching rate of C. sinica larvae on A. gossypii were studied in laboratory. Predating numbers of C. sinica each instar larva increased with the density increase of A. gossypii in different cotton varieties. Most of the predating numbers of C. sinica feeding on A. gossypii in transgenic cotton were higher than those in conventional cotton. Predatory functional response of C. sinica each instar larva was consistent with Holling Ⅱ model pattern. According to the predatory functional response equation, we found that the daily maximum predation of C. sinica feeding on A. gossypii in transgenic cotton were higher than those in conventional cotton, while the treating time of C. sinica feeding on A. gossypii in transgenic cotton were less than those in conventional cotton. The daily maximum predation of C. sinica2nd and3rd instar larva feeding on A. gossypii in SGK321was29%and41%higher than the control. Search rate of C. sinica of each instar larva feeding on A. gossypii in transgenic cotton were higher than those in conventional cotton, and intraspecific interference of C. sinica of each instar larva feeding on A. gossypii in transgenic cotton were stronger than the control. Intensity of scrambling competition of C. sinica1st,2nd and3rd instar larva feeding on A. gossypii in GK12,33B, SGK321was29.6%,27.4%and69.6%, higher than those feeding on A. gossypii in Simian3. Therefore, the transgenic cotton had no harmful effects to the predation of C. sinica larva.
4. In the food chain of transgenic cotton-A. gossypii-honeydew-Messor aciculatus and transgenic cotton-A. gossypii-C. sinica larva, we found there were a very high Bt toxin protein concentration of more than1000ng/g in transgenic cotton leaves, while a relatively low concentration in phloem sap compared with that in cotton leaves, which was only0.36% and0.37%at the seedling stage and the flowering stage, respectively. We also detected the Bt toxin protein of a small content in the cotton aphids fed with transgenic cottons, however, this content became a bit larger (1.16times) in A. gossypii feeding on transgenic cottons continuously for multi-generations, which indicats that with the increase of generation,Bt toxin protein accumulated in A. gossypii. There were also remarkable differences in the amount of honeydew secreted by A. gossypii fed with various kinds of transgenic cottons, in the conventional cotton field, the honeydew secreted was4.1g/d, which was2.28times that of SGK321.Bt toxin protein was detected in honeydew secreted by A. gossypii fed with transgenic cottons, but there was no obvious difference between different treatments. A small amount of Bt toxalbumin was also detected in M. aciculatus around the A. gossypii. There was no Bt toxin protein detected in C. sinica larvae of first or second generation feeding on A. gossypii in each transgenic cotton varieties.
1.转基因抗虫棉Bt毒蛋白表达量的时空变化研究结果表明：Bt基因在所有转基因棉花器官中均有表达,不同生育期功能叶中的Bt毒蛋白含量差异显著,变化趋势为先高后低再略有回升,即苗期>蕾期>花期>吐絮期>铃期；蕾、花、铃各部位中的Bt毒蛋白含量的时间变化也有相同趋势。Bt毒蛋白在各时期的含量均表现为功能叶中最高,大于花、蕾等繁殖器官。不同叶位对Bt毒蛋白含量有直接影响。不同花器官中Bt毒蛋白含量有明显差异,表现为花瓣>花蕊>子房>苞片,其中苞片中Bt毒蛋白含量远远低于花瓣等其它花器官。GK12苞片中Bt毒蛋白含量最低,仅为147.9ng/g;最高Bt毒蛋白含量在SGK321功能叶中被检测到,达1630.1ng/g,为GK12苞片中含量的11.0倍。转基因棉叶中杀虫蛋白在土壤环境中的降解趋势表现为初期降解非常迅速,之后趋于平缓。试验开始的第一个月里(2010年9月～2010年10月),GK12、33B、SGK321三个品种中的杀虫蛋白分别降解了76.78%,82.03%和84.26%,至次年4月份,几乎检测不到Bt蛋白,到5月份已完全检测不到Bt蛋白存在。
2.取食不同品种棉花的蚜虫对中华草蛉生长发育及繁殖的影响结果指出：中华草蛉捕食不同品种棉花上的棉蚜后,其死亡率、羽化率以及茧的发育历期、茧重和成虫性比在棉花品种间大多无显著差异；取食SGK321棉田棉蚜的草蛉幼虫1代死亡率为8.89%,还极显著低于对照处理(40.0%),2代死亡率为16.09%,也显著低于对照处理(75.9%)；取食SGK321棉田棉蚜的草蛉幼虫1、2代羽化率较高,分别为对照品种的1.18倍和1.17倍。各处理间草蛉成虫的产卵前期无显著差异；产卵期表现为取食转基因棉田棉蚜的草蛉都显著长于对照处理,其中取食SGK321棉田棉蚜的草蛉1代产卵历期达33.01天,显著高于对照；用取食转基因棉的棉蚜饲喂草蛉,后者的产卵量高于对应处理。其中,取食转基因棉田棉蚜的草蛉1、2代单雌产卵量部分显著高于对照,且取食SGK321棉田棉蚜的草蛉幼虫1代单雌产卵量高达882粒,显著高于对照处理。各处理间草蛉卵的孵化率无显著差异。一系列研究结果均表明,转Bt基因抗虫棉对中华草蛉的生长发育和繁殖无不良影响。取食转基因棉,尤其是SGK321品种上的棉蚜对中华草蛉的生长发育与繁殖似有一定的促进作用。
3.中华草蛉对不同品种棉田棉蚜的功能反应与搜寻效应结果发现,草蛉幼虫取食蚜虫的数量随猎物密度的增加而增大,虽然草蛉对转基因棉田蚜虫的捕食量大多高于常规棉田,但差异不显著。草蛉幼虫各虫龄对棉蚜的捕食功能反应均符合Holling Ⅱ型。功能反应方程指出,草蛉幼虫的最大日捕食量常规棉略低于转基因棉,捕食转基因棉田棉蚜的处理时间短于捕食常规棉田的棉蚜。草蛉2龄、3龄幼虫对SGK321棉田棉蚜的日平均捕食量高于对照29%和41%。研究指出,草蛉各龄幼虫对常规棉田棉蚜的捕食作用率要低于转基因棉处理。草蛉各龄幼虫捕食转基因棉田棉蚜的种内干扰强度高于对照常规棉处理。分摊竞争强度研究结果表明：中华草蛉对转基因棉田棉蚜的分摊竞争强度大于常规棉田,1龄、2龄、3龄中华草蛉对转基因棉田棉蚜的最高分摊竞争强度分别高于对照品种棉田棉蚜的29.6%,27.4%和69.6%。由此可见,转基因棉对中华草蛉的捕食作用无不良影响。
4.Bt毒蛋白在棉花-棉蚜-蜜露-针毛收获蚁中的转移及在棉花-棉蚜-中华草蛉食物链中的传递结果发现：转基因棉叶中存在高浓度的Bt毒蛋白,三个转基因棉品种苗期功能叶的Bt毒蛋含量均达1000ng/g以上,而韧皮部汁液中的Bt毒蛋白含量非常低,苗期、花铃期分别仅为棉叶含量的0.36%和0.37%。取食转基因棉花的棉蚜体内也检测到了Bt毒蛋白,但含量甚微。连续、多代取食转基因棉的棉蚜,其体内Bt毒蛋白含量较高,是取食转基因棉一代棉蚜的1.16倍。表明随着取食代数的增加,毒蛋白在棉蚜体内似有一定的累积。取食各转基因棉的棉蚜分泌的蜜露量差异显著,常规棉田棉蚜泌蜜量达4.1g/d,是SGK321转基因棉棉蚜泌蜜量的2.28倍。取食转基因棉的棉蚜分泌的蜜露中检测到了Bt毒蛋白,但处理间无显著差异。围绕棉蚜的针毛收获蚁体内检测到微量的Bt毒蛋白。1代、2代中华草蛉幼虫捕食不同品种转基因棉田的棉蚜后,其体内均检测不出Bt毒蛋白,表明Bt毒蛋白不经过棉蚜传至其天敌中华草蛉体内。
Transgenic Bt cotton varieties have been commercialized in a large scale. In this study, transgenic cotton varieties (GK12, Xinmian33B, SGK321) and their corresponding conventional cotton controls (Simian3, Xinmian33and Shiyuan321) were used to determine Bt toxin content at different developmental stages and different parts of transgenic cotton. The effects of Aphis gossypii feeding in transgenic cotton on the growth, development, the reproduction, the predation functional responses and search effects of Chrysoperla sinica were evaluated. Transfer of Bt toxin protein among transgenic cotton-A. gossypii-honeydew and nature enemies was studied. The main results were as follows:
1. Temporal and spatial variation of the Bt toxin protein content in transgenic cotton showed that Bt gene was expressed in all organs, and there were significant differences in different growth stages of functional leaves. The variation trend was that the Bt expression was high and then decreased, and rose up a little at last, and seedling stage> bud stage> flowering> boll opening period>boll stage; the temporal variation of the Bt toxin protein content of bud, flower, bell also had similar trends. The spatial dynamics of Bt toxin proteins in the same period were consistent. The Bt toxin protein content of functional leaves was highest, and it was more than flowers, buds and other reproductive organs. Different leaf positions had a direct impact on the protein content of the Bt toxin. Bt toxin protein were significantly different in the different floral organs, that is, petal> flower> ovary> bract, and the Bt toxin protein in the bracts was far below the petals and other floral organs. Bt toxin protein in GK12bracts was the lowest, only147.9ng/g; highest Bt toxin protein was detected in SGK321functional leaves,1630.1ng/g,11.0times as the minimum content. In natural environment, the insecticidal protein in Bt cotton degraded much rapidly in the initial period and then slowered, it reached76.78%,82.03%and84.26%in first month (2010.9～2010.10) respectively in GK12, Xinmian33B, SGK321, and undetectable in late May.
2. Impact on the growth development and the reproduction of C. sinica feeding on A. gossypii in different varieties of cotton indicated that after preying on the A. gossypii on different varieties of cotton, the mortality rate, the emergence rate and cocoon developmental duration, cocoon weight and adult sex ratio of C. sinica in cotton varieties were almost no significant differences; only the mortality rate of the first generation of C. sinica larvae feeding on A. gossypii in SGK321was only8.89%, extremely lower than the control (40.0%). The mortality rate of the second generation was16.09%, significantly lower than the control (75.9%); and the emergence rate of the first and second generations of the C. sinica larvae feeding on the A. gossypii of SGK321was high,1.18and1.17times as the control. There was no significant difference among the treatments of C. sinica adult in preoviposition; The spawning period for C. sinica feeding on the A. gossypii of transgenic cotton were longer than the control; the oviposition duration of the first generation of C. sinica feeding on the A. gossypii in SGK321was up to33.01d, significantly longer than the control. The fecundity of C. sinica feeding on the A. gossypii in transgenic cotton was higher than the control. The fecundity of the first and second generation of the C. sinica feeding on the A. gossypii in33B and SGK321were significantly higher than control, and the fecundity of the first generation of C. sinica feeding on the A. gossypii in SGK321was up to882, which was extremely significantly higher than the control. There was no significant difference in egg hatchability between treatments. A series of studies showed that transgenic cotton had no adverse effects on the growth and reproduction of C. sinica. A. gossypii in transgenic cotton especially SGK321seems to have certain positive role in promoting the growth and development and reproduction of C. sinica.
3. Predation functional response and searching rate of C. sinica larvae on A. gossypii were studied in laboratory. Predating numbers of C. sinica each instar larva increased with the density increase of A. gossypii in different cotton varieties. Most of the predating numbers of C. sinica feeding on A. gossypii in transgenic cotton were higher than those in conventional cotton. Predatory functional response of C. sinica each instar larva was consistent with Holling Ⅱ model pattern. According to the predatory functional response equation, we found that the daily maximum predation of C. sinica feeding on A. gossypii in transgenic cotton were higher than those in conventional cotton, while the treating time of C. sinica feeding on A. gossypii in transgenic cotton were less than those in conventional cotton. The daily maximum predation of C. sinica2nd and3rd instar larva feeding on A. gossypii in SGK321was29%and41%higher than the control. Search rate of C. sinica of each instar larva feeding on A. gossypii in transgenic cotton were higher than those in conventional cotton, and intraspecific interference of C. sinica of each instar larva feeding on A. gossypii in transgenic cotton were stronger than the control. Intensity of scrambling competition of C. sinica1st,2nd and3rd instar larva feeding on A. gossypii in GK12,33B, SGK321was29.6%,27.4%and69.6%, higher than those feeding on A. gossypii in Simian3. Therefore, the transgenic cotton had no harmful effects to the predation of C. sinica larva.
4. In the food chain of transgenic cotton-A. gossypii-honeydew-Messor aciculatus and transgenic cotton-A. gossypii-C. sinica larva, we found there were a very high Bt toxin protein concentration of more than1000ng/g in transgenic cotton leaves, while a relatively low concentration in phloem sap compared with that in cotton leaves, which was only0.36% and0.37%at the seedling stage and the flowering stage, respectively. We also detected the Bt toxin protein of a small content in the cotton aphids fed with transgenic cottons, however, this content became a bit larger (1.16times) in A. gossypii feeding on transgenic cottons continuously for multi-generations, which indicats that with the increase of generation,Bt toxin protein accumulated in A. gossypii. There were also remarkable differences in the amount of honeydew secreted by A. gossypii fed with various kinds of transgenic cottons, in the conventional cotton field, the honeydew secreted was4.1g/d, which was2.28times that of SGK321.Bt toxin protein was detected in honeydew secreted by A. gossypii fed with transgenic cottons, but there was no obvious difference between different treatments. A small amount of Bt toxalbumin was also detected in M. aciculatus around the A. gossypii. There was no Bt toxin protein detected in C. sinica larvae of first or second generation feeding on A. gossypii in each transgenic cotton varieties.
引文
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