艾蒿精油对白纹伊蚊的作用及卵黄发生的影响
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摘要
白纹伊蚊Aedes albopictus Skuse,属于双翅目蚊科,是传播登革热和登革出血热等多种传染病的重要媒介之一。艾篙Artemisia argyi Levl.et Vant为菊科篙属多年生草本植物,艾蒿精油具有很强的杀菌灭虫功效,对人畜低毒,是一种廉价的天然杀虫植物。本文主要研究了艾蒿精油对白纹伊蚊等昆虫的多种生物活性,以及对白纹伊蚊产卵量、卵巢发育、卵黄发生及酶系的影响,为艾蒿精油应用于害虫防治、探讨艾蒿精油对白纹伊蚊卵黄发生的作用机理提供理论基础和依据。主要结果如下:
1.艾蒿精油对白纹伊蚊等昆虫的作用
艾蒿精油对昆虫的作用方式主要有熏蒸、触杀、忌避和杀卵作用,且对不同种类的昆虫有选择性。艾蒿精油对多种昆虫表现出很高的熏蒸毒力,对白纹伊蚊成虫熏蒸0.5h,1h和4h后的LC50分别为0.69,0.59和0.47μL/L;40gL剂量下熏蒸96h对玉米象和赤拟谷盗成虫的致死率分别为100%和76.3%。艾蒿精油对白纹伊蚊成虫和幼虫的LD_(50)分别为67.6和45.4μg/mL;对赤拟谷盗和玉米象的LD_(50)分别为24.1和39.8μg/mL。艾蒿精油对白纹伊蚊成虫的最长保护时间为5.2小时:而赤拟谷盗成虫与玉米象成虫相比,艾蒿精油对赤拟谷盗的忌避效果更好。艾蒿精油对白纹伊蚊有很好的杀卵作用,各浓度处理组与对照组都存在显著差异。经LD_(10)和LD_(15)处理后白纹伊蚊的产卵量与对照呈显著差异,LD_(15)处理组的孵化率与对照组差异显著。
2.艾蒿精油处理对白纹伊蚊卵巢发育的影响
艾蒿精油处理后对白纹伊蚊的卵子和卵母细胞生长无明显影响,对照组和处理组都经历了前期平稳增长和后期快速增长直至稳定的发育过程,卵子和卵母细胞在各发育阶段的长度均与对照无显著差异。对照组和处理组中卵黄物质在卵母细胞中的沉积都经历了前期快速增长和后期缓慢增长直至稳定的过程,不同的是,从吸血后3小时至33小时,处理组中卵黄物质在卵母细胞中的比例显著低于对照组,在其它发育阶段则差异不显著。对比对照组和处理组的白纹伊蚊卵巢发育的显微结构特点,发现,在卵巢发育前期,处理组卵母细胞中存在较大颗粒的脂滴,对照组中不存在;在对照组中,卵黄物质的沉积起始于吸血后6小时,而处理组起始于吸血后12小时;对照组的卵黄膜形成在吸血后36小时,而处理组在吸血后48小时;在吸血后72小时,对照组中的卵壳已经形成,而处理组中还未能形成。精油处理后对卵巢发育超微结构的影响表现在:卵母细胞核内异染色质堆积呈不规则形状;滤泡细胞排列松散,开放间隙异常增大;卵母细胞中卵黄颗粒失去了原来的圆形或椭圆形,呈不规则形状:卵黄膜由多层颗粒排列组成,颗粒间不紧密,有较大缝隙;滋养细胞有死亡的现象,表现在滋养细胞失去了细胞原有的形态,细胞质发生溶解,里面的物质有丢失的情况等。
3.白纹伊蚊卵黄蛋白的纯化及组织定位分析
采用凝胶过滤层析和离子交换层析的方法对白纹伊蚊的卵黄蛋白(Vt)进行了纯化。SDS-PAGE分析表明,白纹伊蚊Vt由大220 KDa和66KDa两个亚基组成。Western免疫印迹结果表明,这两个亚基均能与Vt的多克隆抗体起特异性反应,但与雄虫匀浆液无反应,可以用于Vg/Vt的定性或定量检测。SDS-PAGE及对应Western免疫印迹分析表明,吸血后48小时雌蚊的脂肪体、血淋巴和卵的粗提物中均存在Vt的两个亚基,而雄蚊的血淋巴中不存在。
4.艾蒿精油处理对卵黄发生的影响
采用免疫大白兔的方法,制备得到抗白纹伊蚊Vt的多克隆抗血清,其效价达到1:12800。纯化后的多克隆抗体的重链和轻链的分子量分别为67.9 KDa和43.5KDa。通过ELISA检测发现,与对照组相比,艾蒿精油处理后对白纹伊蚊脂肪体、血淋巴及卵巢中Vg/Vt的发育趋势没有明显的影响。在对照组中,脂肪体中Vg含量的峰值出现在吸血后24h,处理组中的峰值则出现在吸血后36小时,且峰值之间呈现显著差异。对照组血淋巴中Vg含量的峰值都出现在吸血后24h,而处理组的峰值则出现在吸血后36小时,且峰值之间呈现显著差异。在对照组中,卵巢Vt含量在吸血后24小时达到峰值,而处理组则延迟到吸血后48h,但含量没有差异。
5.艾蒿精油处理对白纹伊蚊体内酶系的影响
艾蒿精油处理后,白纹伊蚊体内谷胱甘肽S.转移酶活性被显著激活,处理后36h、48h和72h的比活力分别比对照升高91%、87%和44%,而在处理初期(12h和24h)与对照无显著差异。艾蒿精油处理对酸性磷酸酶和Na~+ -K~+ -ATP酶活性有强烈的抑制作用,处理组都与对照存在显著差异。对碱性磷酸酶的活性表现出先抑制(12h和24h)后激活(36h和48h),72h后又被抑制。
The mosquito,Aedes albopictus Skuse(Diptera:Culicidae) is one of the important intermediary agents who spreads various infectious diseases such as dengue fever and dengue haemorrhagic fever.Artemisia argyi Levl.et Vant(Compositae:Artemisia) is one kind of perennial herb,the essential oil of A.argyi has strong bactericidal and insecticidal effect to insect,but low toxicity to human and livestock,so it is one kind of cheap natural insecticidal plant.In this study,the repellent activity and toxicity of A.argyi oil to A. albopictus,Tribolium castaneum and Sitophilus zeamaisewere studied.Also,its effect on egg laying,ovary development,vitellogenesis and enzymes of adults of A.albopictus were analysed,for the purpose of providing theoretical principle and basis for pest controlling and exploring the mechanism of A.argyi oil on vitellogenesis of A.albopictus. The results were as follows:
1.The effect of A.argyi to insects such as A.albopictus
The action modes of A.argyi oil to insect were fumigant,contact toxicity,repellent and ovicidal activity,and the action is selective to different kinds of insect.A.argyi oil expressed high fumigant toxicity to many kinds of insects.The LC_(50) values(0.5h,1h and 4h) to adult of A.albopictus were 0.69,0.59 and 0.47μL/L;the lethality rates(96h) of adults of T.castaneum and S.zeamaise treated with 40μL were 100%and 76.3%.Also,the LD_(50) values of A.argyi oil against adults and larvae of A.albopictus were 67.6 and 45.4μg/mL;the LD_(50) values were 24.1 and 39.8μg/mL to adults of T.castaneum and S.zeamaise.It was also repellent to the adults of A.albopictus,the longest protection time was 5.2h;T.castaneum was more susceptible than S.zeamaise based on the repellency effect.A.argyi oil had the strong ovicidal activity;all the treatments were significantly different compared with the control.The number of egg laying by A.albopictus was reduced treated with LC_(10) and LC_(15),and the hatchability was inhibited treated with LC_(15) significantly different compared with that of the control.
2.The effect of A.argyi essential oil to the ovary development of A.albopictus
A.argyi oil treatment had no obvious effect on eggs and oocytes development;Both the control group and treatment group experienced a steady growth in earlier stage and a rapid growth in later stage until stability,there was no significant difference between control groups and treated groups on the lengths of the egg and oocyte in each developmental stage.The yolk protein in the oocytes in controls and treatments experienced a rapid growth in earlier stage and a steady growth in later stage until stability,what is the difference is the proportion of yolk protein in the oocytes in treatments was significantly lower than that of the control during 3h to 33 h period ot time after blood-sucking;there was no significant difference in other developmental stages. Compared the microstructure of ovarian development characteristics of A.albopictus of control group with that of treated group,we found that there has large lipid droplets in the oocytes in treatments in earlier developmental stage where the control did not exist;we also found in the control group,the deposition of yolk protein began at 6h after blood meal,whereas the treatment group began at 12h;in the control group,the vitelline membrane formed at 36h after blood meal,whereas at 48h in the treatment group;we also found in the control,the chorion was formed at 72h after blood-sucking,whereas it was not observed in the treatment.The effect on the ultrastructure of A.albopictus ovarian eggs after treated with A.argyi oil was:the heterochromatin in the oocyte nucleus was in irregular shape;follicular cell arranged loose and the opening space increased abnormally; the yolk granules lost its original circle shaped or oval,was in irregular shape in the oocytes;the vitelline membrane was formed by muti-layer of granules,which arranged not closely;there were nurse cells in the egg,which characterized by losing the original form,losing materials and cytoplasm dissolution and so on.
3.The purification of yolk protein of A.albopictus and analysis of tissue localization
Gel filtration chromatography and ion-exchange chromatography were used to purify the vitellin of A.albopictus.SDS-PAGE analysis showed that the vitellin was composed of two subsuits,the large one 220 KDa and the small one 66 KDa.Western-blotting results showed that both of these two subunits acted specific reaction with polyclonal antibody,however,has no action with the homogenate of male.SDS-PAGE and corresponding Western-blotting anasysis showed the two subsuits existed in the fat body and hemolymph of female that has sucked blood for 48 h and its egg extract,but not in hemolymph of male of A.albopictus.
4.The effect of A.argyi essential oil to the vitellogenesis
A polyclonal antiserum against vitellin of A.albopictus was prepared in male rabbits, and the titer reached 1:12 800.We found the heavy chain and light chain of the polyclonal antibody were 67.9KDa and 43.5KDa,respectively.Detected through ELISA, we found that the Vg/Vt development trend in fat body,hemolymph and ovarian in treatment had not been impact when compared with the control.The results showed that in the control group,the Vg content in the fat body achieved peak at 24h after blood-sucking,while the peak appeared at 36h in the treatment group,whereas the two peaks content were significantly defferent;the Vg content peak appeared at 24h after blood-sucking in the control while at 36h in the treatment,and there was no significant difference between them;in the control group,the peak of Vt content was occurred at 24h after blood-sucking while the time was postponed until 48h in the treated group,but no difference between them.
5.The effect of A.argyi essential oil to the enzyme system of A.albopictus
The results showed that after treated with the A.argyi oil,the activity of GST was significantly activated;the activity of GST increased 91%、87%and 44%at 36h,48h and 72h respectively more than that of control,while there was no significant difference at the earlier stage(12h and 24h).A.argyi oil treatment had a strong inhibitory effect on the activities of ACP and Na~+-K~+-ATP,while there were significant differences between all the treatments and that of the related control.The activity of AKP reduced on the first (12h and 24h),then enhanced(36h and 48h),and then inhibited at 72h after blood feeding after treated with A.argyi oil.
1.艾蒿精油对白纹伊蚊等昆虫的作用
艾蒿精油对昆虫的作用方式主要有熏蒸、触杀、忌避和杀卵作用,且对不同种类的昆虫有选择性。艾蒿精油对多种昆虫表现出很高的熏蒸毒力,对白纹伊蚊成虫熏蒸0.5h,1h和4h后的LC50分别为0.69,0.59和0.47μL/L;40gL剂量下熏蒸96h对玉米象和赤拟谷盗成虫的致死率分别为100%和76.3%。艾蒿精油对白纹伊蚊成虫和幼虫的LD_(50)分别为67.6和45.4μg/mL;对赤拟谷盗和玉米象的LD_(50)分别为24.1和39.8μg/mL。艾蒿精油对白纹伊蚊成虫的最长保护时间为5.2小时:而赤拟谷盗成虫与玉米象成虫相比,艾蒿精油对赤拟谷盗的忌避效果更好。艾蒿精油对白纹伊蚊有很好的杀卵作用,各浓度处理组与对照组都存在显著差异。经LD_(10)和LD_(15)处理后白纹伊蚊的产卵量与对照呈显著差异,LD_(15)处理组的孵化率与对照组差异显著。
2.艾蒿精油处理对白纹伊蚊卵巢发育的影响
艾蒿精油处理后对白纹伊蚊的卵子和卵母细胞生长无明显影响,对照组和处理组都经历了前期平稳增长和后期快速增长直至稳定的发育过程,卵子和卵母细胞在各发育阶段的长度均与对照无显著差异。对照组和处理组中卵黄物质在卵母细胞中的沉积都经历了前期快速增长和后期缓慢增长直至稳定的过程,不同的是,从吸血后3小时至33小时,处理组中卵黄物质在卵母细胞中的比例显著低于对照组,在其它发育阶段则差异不显著。对比对照组和处理组的白纹伊蚊卵巢发育的显微结构特点,发现,在卵巢发育前期,处理组卵母细胞中存在较大颗粒的脂滴,对照组中不存在;在对照组中,卵黄物质的沉积起始于吸血后6小时,而处理组起始于吸血后12小时;对照组的卵黄膜形成在吸血后36小时,而处理组在吸血后48小时;在吸血后72小时,对照组中的卵壳已经形成,而处理组中还未能形成。精油处理后对卵巢发育超微结构的影响表现在:卵母细胞核内异染色质堆积呈不规则形状;滤泡细胞排列松散,开放间隙异常增大;卵母细胞中卵黄颗粒失去了原来的圆形或椭圆形,呈不规则形状:卵黄膜由多层颗粒排列组成,颗粒间不紧密,有较大缝隙;滋养细胞有死亡的现象,表现在滋养细胞失去了细胞原有的形态,细胞质发生溶解,里面的物质有丢失的情况等。
3.白纹伊蚊卵黄蛋白的纯化及组织定位分析
采用凝胶过滤层析和离子交换层析的方法对白纹伊蚊的卵黄蛋白(Vt)进行了纯化。SDS-PAGE分析表明,白纹伊蚊Vt由大220 KDa和66KDa两个亚基组成。Western免疫印迹结果表明,这两个亚基均能与Vt的多克隆抗体起特异性反应,但与雄虫匀浆液无反应,可以用于Vg/Vt的定性或定量检测。SDS-PAGE及对应Western免疫印迹分析表明,吸血后48小时雌蚊的脂肪体、血淋巴和卵的粗提物中均存在Vt的两个亚基,而雄蚊的血淋巴中不存在。
4.艾蒿精油处理对卵黄发生的影响
采用免疫大白兔的方法,制备得到抗白纹伊蚊Vt的多克隆抗血清,其效价达到1:12800。纯化后的多克隆抗体的重链和轻链的分子量分别为67.9 KDa和43.5KDa。通过ELISA检测发现,与对照组相比,艾蒿精油处理后对白纹伊蚊脂肪体、血淋巴及卵巢中Vg/Vt的发育趋势没有明显的影响。在对照组中,脂肪体中Vg含量的峰值出现在吸血后24h,处理组中的峰值则出现在吸血后36小时,且峰值之间呈现显著差异。对照组血淋巴中Vg含量的峰值都出现在吸血后24h,而处理组的峰值则出现在吸血后36小时,且峰值之间呈现显著差异。在对照组中,卵巢Vt含量在吸血后24小时达到峰值,而处理组则延迟到吸血后48h,但含量没有差异。
5.艾蒿精油处理对白纹伊蚊体内酶系的影响
艾蒿精油处理后,白纹伊蚊体内谷胱甘肽S.转移酶活性被显著激活,处理后36h、48h和72h的比活力分别比对照升高91%、87%和44%,而在处理初期(12h和24h)与对照无显著差异。艾蒿精油处理对酸性磷酸酶和Na~+ -K~+ -ATP酶活性有强烈的抑制作用,处理组都与对照存在显著差异。对碱性磷酸酶的活性表现出先抑制(12h和24h)后激活(36h和48h),72h后又被抑制。
The mosquito,Aedes albopictus Skuse(Diptera:Culicidae) is one of the important intermediary agents who spreads various infectious diseases such as dengue fever and dengue haemorrhagic fever.Artemisia argyi Levl.et Vant(Compositae:Artemisia) is one kind of perennial herb,the essential oil of A.argyi has strong bactericidal and insecticidal effect to insect,but low toxicity to human and livestock,so it is one kind of cheap natural insecticidal plant.In this study,the repellent activity and toxicity of A.argyi oil to A. albopictus,Tribolium castaneum and Sitophilus zeamaisewere studied.Also,its effect on egg laying,ovary development,vitellogenesis and enzymes of adults of A.albopictus were analysed,for the purpose of providing theoretical principle and basis for pest controlling and exploring the mechanism of A.argyi oil on vitellogenesis of A.albopictus. The results were as follows:
1.The effect of A.argyi to insects such as A.albopictus
The action modes of A.argyi oil to insect were fumigant,contact toxicity,repellent and ovicidal activity,and the action is selective to different kinds of insect.A.argyi oil expressed high fumigant toxicity to many kinds of insects.The LC_(50) values(0.5h,1h and 4h) to adult of A.albopictus were 0.69,0.59 and 0.47μL/L;the lethality rates(96h) of adults of T.castaneum and S.zeamaise treated with 40μL were 100%and 76.3%.Also,the LD_(50) values of A.argyi oil against adults and larvae of A.albopictus were 67.6 and 45.4μg/mL;the LD_(50) values were 24.1 and 39.8μg/mL to adults of T.castaneum and S.zeamaise.It was also repellent to the adults of A.albopictus,the longest protection time was 5.2h;T.castaneum was more susceptible than S.zeamaise based on the repellency effect.A.argyi oil had the strong ovicidal activity;all the treatments were significantly different compared with the control.The number of egg laying by A.albopictus was reduced treated with LC_(10) and LC_(15),and the hatchability was inhibited treated with LC_(15) significantly different compared with that of the control.
2.The effect of A.argyi essential oil to the ovary development of A.albopictus
A.argyi oil treatment had no obvious effect on eggs and oocytes development;Both the control group and treatment group experienced a steady growth in earlier stage and a rapid growth in later stage until stability,there was no significant difference between control groups and treated groups on the lengths of the egg and oocyte in each developmental stage.The yolk protein in the oocytes in controls and treatments experienced a rapid growth in earlier stage and a steady growth in later stage until stability,what is the difference is the proportion of yolk protein in the oocytes in treatments was significantly lower than that of the control during 3h to 33 h period ot time after blood-sucking;there was no significant difference in other developmental stages. Compared the microstructure of ovarian development characteristics of A.albopictus of control group with that of treated group,we found that there has large lipid droplets in the oocytes in treatments in earlier developmental stage where the control did not exist;we also found in the control group,the deposition of yolk protein began at 6h after blood meal,whereas the treatment group began at 12h;in the control group,the vitelline membrane formed at 36h after blood meal,whereas at 48h in the treatment group;we also found in the control,the chorion was formed at 72h after blood-sucking,whereas it was not observed in the treatment.The effect on the ultrastructure of A.albopictus ovarian eggs after treated with A.argyi oil was:the heterochromatin in the oocyte nucleus was in irregular shape;follicular cell arranged loose and the opening space increased abnormally; the yolk granules lost its original circle shaped or oval,was in irregular shape in the oocytes;the vitelline membrane was formed by muti-layer of granules,which arranged not closely;there were nurse cells in the egg,which characterized by losing the original form,losing materials and cytoplasm dissolution and so on.
3.The purification of yolk protein of A.albopictus and analysis of tissue localization
Gel filtration chromatography and ion-exchange chromatography were used to purify the vitellin of A.albopictus.SDS-PAGE analysis showed that the vitellin was composed of two subsuits,the large one 220 KDa and the small one 66 KDa.Western-blotting results showed that both of these two subunits acted specific reaction with polyclonal antibody,however,has no action with the homogenate of male.SDS-PAGE and corresponding Western-blotting anasysis showed the two subsuits existed in the fat body and hemolymph of female that has sucked blood for 48 h and its egg extract,but not in hemolymph of male of A.albopictus.
4.The effect of A.argyi essential oil to the vitellogenesis
A polyclonal antiserum against vitellin of A.albopictus was prepared in male rabbits, and the titer reached 1:12 800.We found the heavy chain and light chain of the polyclonal antibody were 67.9KDa and 43.5KDa,respectively.Detected through ELISA, we found that the Vg/Vt development trend in fat body,hemolymph and ovarian in treatment had not been impact when compared with the control.The results showed that in the control group,the Vg content in the fat body achieved peak at 24h after blood-sucking,while the peak appeared at 36h in the treatment group,whereas the two peaks content were significantly defferent;the Vg content peak appeared at 24h after blood-sucking in the control while at 36h in the treatment,and there was no significant difference between them;in the control group,the peak of Vt content was occurred at 24h after blood-sucking while the time was postponed until 48h in the treated group,but no difference between them.
5.The effect of A.argyi essential oil to the enzyme system of A.albopictus
The results showed that after treated with the A.argyi oil,the activity of GST was significantly activated;the activity of GST increased 91%、87%and 44%at 36h,48h and 72h respectively more than that of control,while there was no significant difference at the earlier stage(12h and 24h).A.argyi oil treatment had a strong inhibitory effect on the activities of ACP and Na~+-K~+-ATP,while there were significant differences between all the treatments and that of the related control.The activity of AKP reduced on the first (12h and 24h),then enhanced(36h and 48h),and then inhibited at 72h after blood feeding after treated with A.argyi oil.
引文
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