赤眼蜂种间Wolbachia转染与生防潜力研究
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摘要
赤眼蜂(Trichogramma spp.)是全世界害虫生物防治中研究最多、应用最广的一类卵寄生性天敌,广泛用于防治玉米、水稻、果树和森林等植物害虫,取得了显著的经济效益和生态效益。节肢动物胞内共生菌沃尔巴克氏体(Wolbachia)由于调控宿主进化和影响宿主生殖等行为以及作为外源基因的载体等领域的应用前景,近年来引起国内外学者的广泛关注。Wolbachia广泛分布于赤眼蜂属寄生蜂中,能诱导赤眼蜂产雌孤雌生殖,且可在种内垂直传播和种间水平传播,已报道有21种赤眼蜂感染Wolbachia。由于Wolbachia具有诱导寄生蜂产雌孤雌生殖与增强雌性生殖力的特性,因而Wolbachia可以作为特定的遗传工具对其宿主进行遗传调控,利用其来改变天敌昆虫的生殖特性,通过将不具产雌孤雌生殖特性的优良天敌蜂种人为感染Wolbachia而诱导寄生蜂行产雌孤雌生殖,从而增加雌蜂比例,降低繁蜂成本,提高生物防治效能具有重要意义。
本论文按Wolbachia供体蜂的筛选;摸索Wolbachia水平人工转染方法并探讨转染机制;转染成功致使生殖方式改变的赤眼蜂生物学特性研究,包括外界环境因子的应答能力,嗅觉反应和种群参数;以及Wolbachia在赤眼蜂体内的适合度和田间大面积释放防治玉米螟的效果等内容相继展开。本论文的主要内容及取得的初步成果如下:
1.利用改良的“Y”型嗅觉仪来测定感染Wolbachia的卷蛾赤眼蜂(Trichogramma cacoeciae)、食胚赤眼蜂(Trichogramma embryophagum)和短管赤眼蜂(Trichogramma pretiosum)3种产雌孤雌生殖赤眼蜂对米蛾卵表和雌蛾腹部鳞片正己烷萃取物的趋性反应,评价其搜索寄主能力、寄生和羽化能力;观察了经抗生素处理后Wolbachia在3种赤眼蜂体内的稳定性,探讨Wolbachia菌被去除后宿主赤眼蜂的生殖能力变化情况。
通过试验发现抗生素处理后食胚赤眼蜂后代羽化雄性最少,其体内Wolbachia与宿主食胚赤眼蜂共生比较稳定,且该蜂嗅觉反应灵敏,寄生能力强,故认为食胚赤眼蜂体内Wolbachia是水平人工转染较理想株系,适合作Wolbachia水平转染的供体。
2.以产雌孤雌生殖食胚赤眼蜂为供体,两性生殖的松毛虫赤眼蜂Trichogrammadendrolimi、玉米螟赤眼蜂Trichogramma ostriniae、螟黄赤眼蜂Trichogramma chilonis和甘蓝夜蛾赤眼蜂Trichogramma brassicae为受体,通过共享食物源、显微注射、取食菌悬液三种方法研究Wolbachia在赤眼蜂种间水平人工传播。结果表明,通过共享食物源方法由食胚赤眼蜂向松毛虫赤眼蜂的种间水平转染获得成功,受体松毛虫赤眼蜂生殖方式由两性生殖方式转变为产雌孤雌生殖,生殖方式稳定;分子检测产雌孤雌松毛虫赤眼蜂体内Wolbachia菌wsp序列与供体食胚赤眼蜂核苷酸同一性达99%,同属于Wolbachia菌群B大组Dei亚组,而与同组Pip亚组寄主米蛾(Corcyra cephalonica)wsp同一性较远为81%。其它供试赤眼蜂种不能通过该方法实现水平转染。利用显微注射法获得了产两性孤雌生殖甘蓝夜蛾赤眼蜂,松毛虫赤眼蜂部分表现了产雌孤雌生殖。赤眼蜂成虫和幼虫取食菌悬液不能实现种间Wolbachia的水平传播。研究结果除可以筛选获得携带Wolbachia且寄生能力强的优良赤眼蜂蜂种或品系,形成现实生产力外,有望提供新的调控生物生殖方式的手段。
3.以成功转染Wolbachia产雌孤雌生殖松毛虫赤眼蜂为材料,分析了25℃和32℃两种温度下交配处理、不同交配组合及抗生素处理后产雌孤雌生殖松毛虫赤眼蜂的寄生数和羽化数等生物学指标。结果表明,25℃温度条件下,产雌孤雌赤眼蜂连续多代培养生殖方式稳定,与雄性交配尽管降低寄生能力但生殖方式没有改变:32℃高温条件下,与雄蜂交配和未交配的产雌孤雌松毛虫赤眼蜂连续处理4代(大约30天)体内Wolbachia都完全失活,恢复产雄孤雌生殖方式。实验表明,与雄蜂交配不会改变感染Wolbachia松毛虫赤眼蜂产雌孤雌的生殖方式,抗生素处理和持续至少20天的32℃高温才是改变其生殖方式的决定因素。抗生素可以去除松毛虫赤眼蜂体内的Wolbachia并恢复两性的生殖方式,供试抗生素在中低浓度(25mg/mL)诱导出的雄蜂最多,而且经抗生素去除获得的雄性和雌性具有正常的生殖功能。本研究明确外界生态因子对感染Wolbachia松毛虫赤眼蜂产雌孤雌生殖方式遗传稳定性的影响,为感染Wolbachia松毛虫赤眼蜂的阳间释放提供理论依据。
4.采用与未感染的两性品系松毛虫赤眼蜂相对比的方法,研究了产雌孤雌生殖松毛虫赤眼蜂对不同寄主挥发物的嗅觉反应能力、寄生及羽化能力。结果表明,产雌孤雌品系松毛虫赤眼蜂对供试的几种鳞翅目害虫(米蛾Corcyra cephalonica、玉米螟Ostriniafurnacalis、柞蚕Antheraea pernyi)卵表和雌蛾腹部鳞片挥发物均有不同程度的趋性,对玉米螟卵表挥发物的趋性较两性品系强,在恒定温湿度(25℃,RH70%)条件下对几种寄主卵的寄生能力均强于两性品系松毛虫赤眼蜂,但两者间差异不明显。在不同蜂卵比接蜂的试验中,30粒(1块)玉米螟卵,除接入2头蜂时,正常两性品系的松毛虫赤眼蜂寄生能力高于产雌孤雌品系外,接入其它数量如1、3、4头蜂时产雌孤雌品系寄生能力均高于两性品系松毛虫赤眼蜂。总体来看,产雌孤雌品系松毛虫赤眼蜂在不同蜂卵比情况下,对玉米螟卵的寄生能力要强于正常两性品系。本项研究为感染Wolbachia的松毛虫赤眼蜂田间防治玉米螟及其它重要农业害虫提供了助力。
5.为探讨感染Wolbachia的产雌孤雌松毛虫赤眼蜂的最佳繁殖条件。本试验对感染Wolbachia松毛虫赤眼蜂不同温度下的生殖参数进行了研究。以柞蚕卵为寄主,在16℃、20℃、24℃、28℃和32℃5个恒温条件下松毛虫赤眼蜂发育、存活和繁殖的情况,构建了相应温度下的实验种群生命表。结果表明,温度24℃～28℃为繁殖感染Wolbachia产雌孤雌松毛虫赤眼蜂的理想温度条件,可以作为松毛虫赤眼蜂的工厂化生产作业指标。
6.对人工水平转染Wolbachia产雌孤雌松毛虫赤眼蜂的田间防治效果进行评价。试验选取辽宁沈阳和岫岩两地区,约600亩玉米田连续两年释放产雌孤雌松毛虫赤眼蜂,评价其对玉米螟的防治效果。通过两年调查发现:产雌孤雌和正常两性松毛虫赤眼蜂防治一代玉米螟防效达到40%以上;同时放蜂田玉米每亩鲜重和干重都明显高于对照未放蜂田,两个品系的松毛虫赤眼蜂在防虫效果和玉米增产等指标均没有明显的差异,表明产雌孤雌品系松毛虫赤眼蜂至少可以在大田应用上取得与正常的两性品系松毛虫赤眼蜂同期的防治效果,而且同比应用潜力可能更大。
Trichogramma spp.is the one of the egg parasitoids controlling insect pests to manage many pests from maize,rice,cottons,vegetables and forests wiedly and successfully. Wolbachia,intracellular symbiotic bacteria of arthropods,in recent years had aroused wide attention from scholars because they could regulate biological evolution and reproductive behavior of the host,and present prospects in some applications areas such as a carrier of exogenous gene.Wolbachia could infect the many species of parasitic Trichogramma, resulting in female offspring of thelytokous Trichogramma,and could be transfected vertically and horizontally among species.Wolbachia could induce thelytoky parthenogenesis of their parasitic Trichogramma and enhance female fertility characteristic,so they could be used as specific genetic tools manipulating their host to change their reproductive characteristics.The better parasitic Trichogramma without producing female offsprings could be rebuilt through infection by Wolbachia,and finally Trichogramma increase female rate and reduce costs.It was useful to improve the effectiveness of biological control.In this thesis, firstly,Trichogramma infected by Wolbachia were screened to identify the better donor to transfect other species.Secondly,we try to make horizontal transfection by artificial method and clarify the mechanism of transfection.Thirdly,the biological characteristics of the target Trichogramma with producing female were analyzed included the response capacity to external environmental factors,olfactory responses,population parameters and the fitness of Wolbachia in the Trichogramma.Finally we released Trichogramma to survey the effects to control corn borer in field with a large area.The results are as follows:
The three kinds of thelytokous T.cacoeciae,T.embryophagum and T.pretiosum response to separately by hexane extracts from the eggs surface of Corcyra cephalonica and abdominal scales of female C.cephalonica were measured by using the improved Y-tube olfactometer.The results helped to evaluate the capacibility of Trichogramma's searching its host,parasitic and eclosion.To uncover the role of Wolbachia in regulating the reproductive capacity of Trichogrammas,the stability of Wolbachia in three kinds of Trichogrammas after antibiotic treatment was observed.The results indicated that the eclosion male offspring of T embryophagum are the smallest after antibiotics treatment,the relationship of Wolbachia and their symbiotic host-T.embryophagum was more stable than others.Combining with sensitive olfactory responses and strong parasitic ability,Wolbachia in T.embryophagum were the better strains to make artificial transfection,and used as donor of horizontal transfection.
Taken thelytokous T.embryophagum as donor,gender reproductive T.dendrolimi,T. ostriniae,T.chilonis and T.brassicae as receptor,the horizontal artificial transfection Wolbachia among Trichogramma species was investigated through the three methods involved in sharing of food source,feeding bacteria suspension and microinjection. Wolbachia was transfected successfully from T.embryophagum to T.dendrolimi by the method of sharing of food source,and the reproductive patterns of receptor T.dendrolimi were changed to thelytoky parthenogenesis with genic stability.Furthermore,the identity of wsp sequence from Wolbachia between thelytokous T.dendrolimi and T.embryophagum reached to 99%,both belonging to“Dei”sub-group of B-group in Wolbachia,while identity between thelytokous T.dendrolimi and C.cephalonica that was classified into“Pip”sub-group of B-large group was 81%.However other Trichogrammas could not accept Wolbachia.Under microinjection,gender thelytoky of T.brassicae were transformed to thelytoky parthenogenesis,and T.dendrolimi showed partial thelytoky parthenogenesis. Horizontal transfection of Wolbachia could not be achieved between Trichogramma species by mature and young Trichogramma feeding bacteria suspension.Trichogramma species or lines embracing Wolbachia with better parasitic ability were obtained to create productive forces,and new means to regulate biological reproduction.
The characteristic of host Trichogramma affected by Wolbachia was diversity and uncertainty.So it is necessary to elucidate the impact of the symbiotic bacteria Wolbachia on Trichogramma.The biological indicators of thelytokous T.dendrolimi including parasitic and eclosion number were analyzed under mating treatments,different combination treatments and antibiotic treatments at 25℃and 32℃.The results indicated that under the conditions of 25℃,the reproduction mode of thelytokous Trichogramma cultured with many generations was fixed.After thelytokous Trichogrammas mating with male,reproduction mode not be changed besides Parasitic ability decreased.Under 32℃,whether Trichogramma mating with male or not were cultured four generations continuously(about 30 days),Wolbachia could be inactivate completely and restore the way of thelytoky parthenogenesis.The results showed that reproduction mode of thelytokous T dendrolimi would not be changed for mating with male.The determination factors of changing mode of reproduction were antibiotic treatment and high temperature 32℃for 20 days at least. Antibiotics may remove Wolbachia in T.dendrolimi and restore the reproduction of gender. Most of male Trichogramma were present exposered antibiotics with middle-low concentration(25mg/mL).The impacts of ecological factors on the genetic stability of thelytokous T.dendrolimi were showed,and theoretical basis were provided releasing T. dendrolimi infected by Wolbachia in field.
Olfactory response of thelytokous T.dendrolimi to volatile infochemicals of host, parasitic capacity and eclosion ability under different rate of wasps to eggs were studied.The thelytokous T.dendrolimi could be attracted in varying degrees by the eggs surface of Corcyra cephalonica and volatile infochemicals of abdominal scales of female C. cephalonica in several provided Lepidoptera pests(C.cephalonica,O.furnacalis,and A. pernyi).The T.dendrolimi response to the eggs surface of corn borer was stronger than gender lines.Under constant temperature and humidity(25℃,RH 70%),the parasitic capacity of T.dendrolimi infected by Wolbachia to several kinds of host eggs was stronger than gender T.dendrolimi's,but the difference was not significant.Focusing on different rate of wasp eggs,the parasitic capacity of thelytokous T.dendrolimi was higher than normal gender lines' when inoculating 1,3 or 4 except 2 of 30(1 piece) corn borer eggs in T. dendrolimi.In summary,the parasitic capacity of thelytokous T.dendrolimi to corn borer eggs was higher than normal gender lines' in different rate of wasp eggs.This research could provide theoretic basis for T.dendrolimi infectted by Wolbachia in controlling corn borer and other important agricultural pests in field.
The reproductive parameters of thelytokous T.dendrolimi infected by Wolbachia were studied at different temperatures in order to investigate the best breeding conditions of them. The biological data including development,surviving rate and reproduction of T.dendrolimi infected by Wolbachia were observed at 16℃,20℃,24℃,28℃and 32℃respectively.The population life table was constructed under the corresponding temperature. The results suggested that the optimal reproduction temperature of thelytokous T.dendrolimi infected by Wolbachia was 24℃-28℃.
Thelytokous T.dendrolimi were released into corn field in the regions of Xiuyan and Shenyang in Liaoning province for two consecutive years,to evaluate effects on controlling corn borer.Through the two-year investigation,it was found that thelytokous T.dendrolimi and gender T.dendrolimi had achieved some effect in controlling corn borer to increase yield more than 40%.Corn fresh weight and dry weight per acre in fields with T.dendrolimi were significantly higher than without T.dendrolimi's.There were not significant differences between the Thelytokous T.dendrolimi and normal T.dendrolimi lines in pest control,corn yield and other indicators.So thelytokous T.dendrolimi in controlling pests was similar to gender T.dendrolimi,and may be greater in potential applications.
本论文按Wolbachia供体蜂的筛选;摸索Wolbachia水平人工转染方法并探讨转染机制;转染成功致使生殖方式改变的赤眼蜂生物学特性研究,包括外界环境因子的应答能力,嗅觉反应和种群参数;以及Wolbachia在赤眼蜂体内的适合度和田间大面积释放防治玉米螟的效果等内容相继展开。本论文的主要内容及取得的初步成果如下:
1.利用改良的“Y”型嗅觉仪来测定感染Wolbachia的卷蛾赤眼蜂(Trichogramma cacoeciae)、食胚赤眼蜂(Trichogramma embryophagum)和短管赤眼蜂(Trichogramma pretiosum)3种产雌孤雌生殖赤眼蜂对米蛾卵表和雌蛾腹部鳞片正己烷萃取物的趋性反应,评价其搜索寄主能力、寄生和羽化能力;观察了经抗生素处理后Wolbachia在3种赤眼蜂体内的稳定性,探讨Wolbachia菌被去除后宿主赤眼蜂的生殖能力变化情况。
通过试验发现抗生素处理后食胚赤眼蜂后代羽化雄性最少,其体内Wolbachia与宿主食胚赤眼蜂共生比较稳定,且该蜂嗅觉反应灵敏,寄生能力强,故认为食胚赤眼蜂体内Wolbachia是水平人工转染较理想株系,适合作Wolbachia水平转染的供体。
2.以产雌孤雌生殖食胚赤眼蜂为供体,两性生殖的松毛虫赤眼蜂Trichogrammadendrolimi、玉米螟赤眼蜂Trichogramma ostriniae、螟黄赤眼蜂Trichogramma chilonis和甘蓝夜蛾赤眼蜂Trichogramma brassicae为受体,通过共享食物源、显微注射、取食菌悬液三种方法研究Wolbachia在赤眼蜂种间水平人工传播。结果表明,通过共享食物源方法由食胚赤眼蜂向松毛虫赤眼蜂的种间水平转染获得成功,受体松毛虫赤眼蜂生殖方式由两性生殖方式转变为产雌孤雌生殖,生殖方式稳定;分子检测产雌孤雌松毛虫赤眼蜂体内Wolbachia菌wsp序列与供体食胚赤眼蜂核苷酸同一性达99%,同属于Wolbachia菌群B大组Dei亚组,而与同组Pip亚组寄主米蛾(Corcyra cephalonica)wsp同一性较远为81%。其它供试赤眼蜂种不能通过该方法实现水平转染。利用显微注射法获得了产两性孤雌生殖甘蓝夜蛾赤眼蜂,松毛虫赤眼蜂部分表现了产雌孤雌生殖。赤眼蜂成虫和幼虫取食菌悬液不能实现种间Wolbachia的水平传播。研究结果除可以筛选获得携带Wolbachia且寄生能力强的优良赤眼蜂蜂种或品系,形成现实生产力外,有望提供新的调控生物生殖方式的手段。
3.以成功转染Wolbachia产雌孤雌生殖松毛虫赤眼蜂为材料,分析了25℃和32℃两种温度下交配处理、不同交配组合及抗生素处理后产雌孤雌生殖松毛虫赤眼蜂的寄生数和羽化数等生物学指标。结果表明,25℃温度条件下,产雌孤雌赤眼蜂连续多代培养生殖方式稳定,与雄性交配尽管降低寄生能力但生殖方式没有改变:32℃高温条件下,与雄蜂交配和未交配的产雌孤雌松毛虫赤眼蜂连续处理4代(大约30天)体内Wolbachia都完全失活,恢复产雄孤雌生殖方式。实验表明,与雄蜂交配不会改变感染Wolbachia松毛虫赤眼蜂产雌孤雌的生殖方式,抗生素处理和持续至少20天的32℃高温才是改变其生殖方式的决定因素。抗生素可以去除松毛虫赤眼蜂体内的Wolbachia并恢复两性的生殖方式,供试抗生素在中低浓度(25mg/mL)诱导出的雄蜂最多,而且经抗生素去除获得的雄性和雌性具有正常的生殖功能。本研究明确外界生态因子对感染Wolbachia松毛虫赤眼蜂产雌孤雌生殖方式遗传稳定性的影响,为感染Wolbachia松毛虫赤眼蜂的阳间释放提供理论依据。
4.采用与未感染的两性品系松毛虫赤眼蜂相对比的方法,研究了产雌孤雌生殖松毛虫赤眼蜂对不同寄主挥发物的嗅觉反应能力、寄生及羽化能力。结果表明,产雌孤雌品系松毛虫赤眼蜂对供试的几种鳞翅目害虫(米蛾Corcyra cephalonica、玉米螟Ostriniafurnacalis、柞蚕Antheraea pernyi)卵表和雌蛾腹部鳞片挥发物均有不同程度的趋性,对玉米螟卵表挥发物的趋性较两性品系强,在恒定温湿度(25℃,RH70%)条件下对几种寄主卵的寄生能力均强于两性品系松毛虫赤眼蜂,但两者间差异不明显。在不同蜂卵比接蜂的试验中,30粒(1块)玉米螟卵,除接入2头蜂时,正常两性品系的松毛虫赤眼蜂寄生能力高于产雌孤雌品系外,接入其它数量如1、3、4头蜂时产雌孤雌品系寄生能力均高于两性品系松毛虫赤眼蜂。总体来看,产雌孤雌品系松毛虫赤眼蜂在不同蜂卵比情况下,对玉米螟卵的寄生能力要强于正常两性品系。本项研究为感染Wolbachia的松毛虫赤眼蜂田间防治玉米螟及其它重要农业害虫提供了助力。
5.为探讨感染Wolbachia的产雌孤雌松毛虫赤眼蜂的最佳繁殖条件。本试验对感染Wolbachia松毛虫赤眼蜂不同温度下的生殖参数进行了研究。以柞蚕卵为寄主,在16℃、20℃、24℃、28℃和32℃5个恒温条件下松毛虫赤眼蜂发育、存活和繁殖的情况,构建了相应温度下的实验种群生命表。结果表明,温度24℃～28℃为繁殖感染Wolbachia产雌孤雌松毛虫赤眼蜂的理想温度条件,可以作为松毛虫赤眼蜂的工厂化生产作业指标。
6.对人工水平转染Wolbachia产雌孤雌松毛虫赤眼蜂的田间防治效果进行评价。试验选取辽宁沈阳和岫岩两地区,约600亩玉米田连续两年释放产雌孤雌松毛虫赤眼蜂,评价其对玉米螟的防治效果。通过两年调查发现:产雌孤雌和正常两性松毛虫赤眼蜂防治一代玉米螟防效达到40%以上;同时放蜂田玉米每亩鲜重和干重都明显高于对照未放蜂田,两个品系的松毛虫赤眼蜂在防虫效果和玉米增产等指标均没有明显的差异,表明产雌孤雌品系松毛虫赤眼蜂至少可以在大田应用上取得与正常的两性品系松毛虫赤眼蜂同期的防治效果,而且同比应用潜力可能更大。
Trichogramma spp.is the one of the egg parasitoids controlling insect pests to manage many pests from maize,rice,cottons,vegetables and forests wiedly and successfully. Wolbachia,intracellular symbiotic bacteria of arthropods,in recent years had aroused wide attention from scholars because they could regulate biological evolution and reproductive behavior of the host,and present prospects in some applications areas such as a carrier of exogenous gene.Wolbachia could infect the many species of parasitic Trichogramma, resulting in female offspring of thelytokous Trichogramma,and could be transfected vertically and horizontally among species.Wolbachia could induce thelytoky parthenogenesis of their parasitic Trichogramma and enhance female fertility characteristic,so they could be used as specific genetic tools manipulating their host to change their reproductive characteristics.The better parasitic Trichogramma without producing female offsprings could be rebuilt through infection by Wolbachia,and finally Trichogramma increase female rate and reduce costs.It was useful to improve the effectiveness of biological control.In this thesis, firstly,Trichogramma infected by Wolbachia were screened to identify the better donor to transfect other species.Secondly,we try to make horizontal transfection by artificial method and clarify the mechanism of transfection.Thirdly,the biological characteristics of the target Trichogramma with producing female were analyzed included the response capacity to external environmental factors,olfactory responses,population parameters and the fitness of Wolbachia in the Trichogramma.Finally we released Trichogramma to survey the effects to control corn borer in field with a large area.The results are as follows:
The three kinds of thelytokous T.cacoeciae,T.embryophagum and T.pretiosum response to separately by hexane extracts from the eggs surface of Corcyra cephalonica and abdominal scales of female C.cephalonica were measured by using the improved Y-tube olfactometer.The results helped to evaluate the capacibility of Trichogramma's searching its host,parasitic and eclosion.To uncover the role of Wolbachia in regulating the reproductive capacity of Trichogrammas,the stability of Wolbachia in three kinds of Trichogrammas after antibiotic treatment was observed.The results indicated that the eclosion male offspring of T embryophagum are the smallest after antibiotics treatment,the relationship of Wolbachia and their symbiotic host-T.embryophagum was more stable than others.Combining with sensitive olfactory responses and strong parasitic ability,Wolbachia in T.embryophagum were the better strains to make artificial transfection,and used as donor of horizontal transfection.
Taken thelytokous T.embryophagum as donor,gender reproductive T.dendrolimi,T. ostriniae,T.chilonis and T.brassicae as receptor,the horizontal artificial transfection Wolbachia among Trichogramma species was investigated through the three methods involved in sharing of food source,feeding bacteria suspension and microinjection. Wolbachia was transfected successfully from T.embryophagum to T.dendrolimi by the method of sharing of food source,and the reproductive patterns of receptor T.dendrolimi were changed to thelytoky parthenogenesis with genic stability.Furthermore,the identity of wsp sequence from Wolbachia between thelytokous T.dendrolimi and T.embryophagum reached to 99%,both belonging to“Dei”sub-group of B-group in Wolbachia,while identity between thelytokous T.dendrolimi and C.cephalonica that was classified into“Pip”sub-group of B-large group was 81%.However other Trichogrammas could not accept Wolbachia.Under microinjection,gender thelytoky of T.brassicae were transformed to thelytoky parthenogenesis,and T.dendrolimi showed partial thelytoky parthenogenesis. Horizontal transfection of Wolbachia could not be achieved between Trichogramma species by mature and young Trichogramma feeding bacteria suspension.Trichogramma species or lines embracing Wolbachia with better parasitic ability were obtained to create productive forces,and new means to regulate biological reproduction.
The characteristic of host Trichogramma affected by Wolbachia was diversity and uncertainty.So it is necessary to elucidate the impact of the symbiotic bacteria Wolbachia on Trichogramma.The biological indicators of thelytokous T.dendrolimi including parasitic and eclosion number were analyzed under mating treatments,different combination treatments and antibiotic treatments at 25℃and 32℃.The results indicated that under the conditions of 25℃,the reproduction mode of thelytokous Trichogramma cultured with many generations was fixed.After thelytokous Trichogrammas mating with male,reproduction mode not be changed besides Parasitic ability decreased.Under 32℃,whether Trichogramma mating with male or not were cultured four generations continuously(about 30 days),Wolbachia could be inactivate completely and restore the way of thelytoky parthenogenesis.The results showed that reproduction mode of thelytokous T dendrolimi would not be changed for mating with male.The determination factors of changing mode of reproduction were antibiotic treatment and high temperature 32℃for 20 days at least. Antibiotics may remove Wolbachia in T.dendrolimi and restore the reproduction of gender. Most of male Trichogramma were present exposered antibiotics with middle-low concentration(25mg/mL).The impacts of ecological factors on the genetic stability of thelytokous T.dendrolimi were showed,and theoretical basis were provided releasing T. dendrolimi infected by Wolbachia in field.
Olfactory response of thelytokous T.dendrolimi to volatile infochemicals of host, parasitic capacity and eclosion ability under different rate of wasps to eggs were studied.The thelytokous T.dendrolimi could be attracted in varying degrees by the eggs surface of Corcyra cephalonica and volatile infochemicals of abdominal scales of female C. cephalonica in several provided Lepidoptera pests(C.cephalonica,O.furnacalis,and A. pernyi).The T.dendrolimi response to the eggs surface of corn borer was stronger than gender lines.Under constant temperature and humidity(25℃,RH 70%),the parasitic capacity of T.dendrolimi infected by Wolbachia to several kinds of host eggs was stronger than gender T.dendrolimi's,but the difference was not significant.Focusing on different rate of wasp eggs,the parasitic capacity of thelytokous T.dendrolimi was higher than normal gender lines' when inoculating 1,3 or 4 except 2 of 30(1 piece) corn borer eggs in T. dendrolimi.In summary,the parasitic capacity of thelytokous T.dendrolimi to corn borer eggs was higher than normal gender lines' in different rate of wasp eggs.This research could provide theoretic basis for T.dendrolimi infectted by Wolbachia in controlling corn borer and other important agricultural pests in field.
The reproductive parameters of thelytokous T.dendrolimi infected by Wolbachia were studied at different temperatures in order to investigate the best breeding conditions of them. The biological data including development,surviving rate and reproduction of T.dendrolimi infected by Wolbachia were observed at 16℃,20℃,24℃,28℃and 32℃respectively.The population life table was constructed under the corresponding temperature. The results suggested that the optimal reproduction temperature of thelytokous T.dendrolimi infected by Wolbachia was 24℃-28℃.
Thelytokous T.dendrolimi were released into corn field in the regions of Xiuyan and Shenyang in Liaoning province for two consecutive years,to evaluate effects on controlling corn borer.Through the two-year investigation,it was found that thelytokous T.dendrolimi and gender T.dendrolimi had achieved some effect in controlling corn borer to increase yield more than 40%.Corn fresh weight and dry weight per acre in fields with T.dendrolimi were significantly higher than without T.dendrolimi's.There were not significant differences between the Thelytokous T.dendrolimi and normal T.dendrolimi lines in pest control,corn yield and other indicators.So thelytokous T.dendrolimi in controlling pests was similar to gender T.dendrolimi,and may be greater in potential applications.
引文
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