斑痣悬茧蜂过寄生行为的研究
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摘要
过寄生是指在1头寄主体内(上)产卵数超出其最大存活数的现象。过寄生不仅给进化生态学提出关于其适应意义的理论问题,而且也是利用天敌昆虫开展生物防治实践面临的重要问题。过寄生在单寄生性昆虫中经常发生,但对其发生的机理却一直存有争议。本学位论文以斑痣悬茧蜂(Meteorus pulchricornis (Wesmae1)为模式生物,主要针对4个科学问题展开研究：i)寄生蜂是否能够辨别出过寄生寄主?ii)过寄生对寄生蟑适合度的不良作用是否随寄主品质不同而变化?iii)影响过寄生发生的潜在因素之间是否存在复杂的互作关系?iv)行为的时间特征是否可用于评价过寄生发生的倾向。论文研究获得以下主要研究结果和结论。
1为探究斑痣悬茧蜂在寄主斑块上的驻留时间与过寄生程度(产卵数)的相关性,采用单选试验设计,观察和分析了斑痣悬茧蜂在斑块内的停留时间与其在寄主幼虫体内产卵数的关系。结果表明,寄生蜂在斑块内驻留的时间影响寄生蜂在每头寄主上的产卵数,1-3h内的产卵平均数分别为0.77、0.9、2.5粒；8h的产卵平均数增加到5.8粒。根据研究推断,在高品质(未被寄生)寄主缺乏的情况下,斑痣悬茧蜂倾向于过寄生。
2为探究寄主品质如何影响过寄生对子代蜂适合度的胁迫作用,采用单选试验设计,研究了以下因素对过寄生行为及其子代发育的影响：不同种寄主(甜菜夜蛾、棉铃虫和斜纹夜蛾),同一种寄主不同龄期寄主幼虫以及取食不同寄主植物。结果表明,过寄生不利于子代蜂的生长发育,导致子代蜂发育历期延长、死亡率增加、体型变小,寿命变短。发育适合度的损耗在不同种类寄主、不同龄期、取食不同植物寄主间发生变化。过寄生甜菜夜蛾幼虫羽化的子代蜂的发育历期、后足胫节比过寄生棉铃虫、斜纹夜羽化的子代蜂的短。与对照不同,过寄生不同龄期的寄主幼虫的子代蜂发育历期、后足胫节、寿命随着龄期的增加而增加；但中间龄期羽化子代蜂的存活率高于从其他龄期羽化的子代蜂。过寄生寄主幼虫取食大豆、棉花和小白菜后的子代蜂发育适合度均显著降低,寄生方式与寄主取食的植物对子代蜂的体型大小有互作效应,与对照相比,从过寄生取食大豆的寄主幼虫中羽化的子代蜂体型减小了22.8%；从过寄生取食棉花的寄主幼虫中羽化的子代蜂体型减小了22.5%；从过寄生取食小白菜的寄主幼虫羽化的子代蜂体型减小了25.0%。据试验研究结果推论,过寄生对寄生蜂适合度的不利影响随寄主品质而变化。
3为探究竞争、寄主充裕程度、卵限制等因素如何影响寄生蜂的过寄生行为,观察了种内竞争、寄主相遇率、寄生蜂抱卵量等对过寄生行为的影响。用基于负二项分布的对数线性模型分析影响过寄生的因素结果表明,种内竞争不影响过寄生发生数量；高抱卵量寄生蜂过寄生寄主的数量高于低抱卵量寄生蜂；寄主相遇率影响过寄生数量,表现为与寄主相遇率高的寄生蜂发生过寄生的次数显著高于与寄主相遇率低的寄生蜂；种内竞争与寄主相遇率对寄生蜂过寄生寄主的数量有互作效应。用识别时间估计寄主被过寄生的风险分析表明,寄主相遇率影响寄主被过寄生的概率,抱卵量和寄主相遇率对过寄生概率具有显著互作影响。据试验结果推论,种内竞争、寄主丰盛度和抱卵量等潜在因素通常互作影响过寄生的发生。
4为评价斑痣悬茧蜂的寄主鉴别能力及其影响因素,采用双选试验研究了寄生蜂的寄生经历和寄主被寄生后的间隔时间对斑痣悬茧蜂在健康寄主和被寄生寄主的选择性。对选择频次进行的分析表明,寄主被寄生后的间隔时间和寄生蜂的寄生经历均对过寄生发生的概率均有显著影响,过寄生概率随寄主被寄生后的间隔时间延长而降低；有寄生经历的寄生蜂发生过寄生的概率低于无寄生经历的寄生蜂。首次测度寄生蜂的寄主识别时间并用COX比例风险模型进行拟合表明,发生过寄生的风险随寄主被寄生后间隔时间的延长而减小,因寄生蜂具有过寄生经历而减小,但寄生蜂具有单寄生经历对过寄生风险没有显著影响。据研究结果推论,斑痣悬茧蜂不仅能够辨别被寄生寄主,而且该能力随寄主被初次寄生后间隔时间延长以及寄生经历而增强。
5野外释放-回收试验的结果表明,斑痣悬茧蜂寄生率为54.2%(168/3l0),过寄生率为23.9%(74/310),寄主体重对寄生蜂在单头寄主上的产卵数有显著影响,随着寄主体重的增加而降低；寄生概率随寄主龄期的增加而降低,但过寄生概率随着寄主龄期的增大而升高；寄生概率随着体重增大而降低,但过寄生概率不受寄主体重影响。据调查结果推论,野外环境中斑痣悬茧蜂可能发生过寄生,而且过寄生程度与产卵时寄主发育时期有关。
6寄生蜂对寄主的识别时间长短反映了其对寄主品质的评价结果。本研究观察测定了斑痣悬茧蜂在评价寄主中所用的识别时间,利用COX比例风险模型拟合识别时间与相关因素的关系,以估计这些因素对过寄生发生倾向的影响。研究结果表明,寄生蜂在决定进行过寄生之前评价寄主所用的识别时间,反映了其对寄主的接受程度,用时长表明寄主质量较差(过寄生),意味着该寄主被寄生的风险较低。研究结果说明,观察寄生蜂在辨别寄主中的识别时间为评价其过寄生行为提供了一个新的特征。
Parasitoid deposits a clutch of eggs (the clutch may consist of a single egg) into a host that has already been parasitized by a member of the same species or itself, a behavior called "superparasitism", and the ensuing larval competition ends in the death of all but one. Superparasitism was not only a focus of theoretical study in evolutionary ecology to understand its adaptation, but also an issue in the practice of biological control using parasitoids to control pests in agriculture. Superparasitism is a common phenomenon in solitary parasitoids, but so far the mechanism of superparasitism is still open to debate. Using Meteorus pulchricornis (Wesmael)(Hymenoptera:Braconidae) as model system, this study was conducted to address four questions:i) whether the parasitoid can discriminate between healthy and parasitized hosts; ii) whether the fitness costs of superparasitism on parasitids vary with host quality; iii) whether potential factors interact to affect superparasitism; iv) whether time-associated behavioral traits might be utilized to evaluate the propensity of superparasitism.The main results and conclusions are summarized as follows.
1A non-choice test was performed to examine the relationship between residence time on host patches and extent of superparasitism as measured by number of eggs per host. The results showed that the residence time had a significant effect on the number of eggs laid in a host. When the residence time was from1to3hours, an average of eggs in a host was0.77,0.9, and2.5respectively, but when the residence time was8hours, an average of5.8eggs was laid in a host. The study suggested that M. pulchricornis have an increasing tendency toward superparasitism with decreasing availability of high-quality hosts.
2In order to examine effects of host quality attributes on fitness consequences of superparasitism for development of parasitoid progeny. A single choice test was used to observe influences of host species, instars, and plant species respectively on the host suitability in superparasitism for the development of parasitoid progeny in a solitary endoparasitoid, M. pulchricornis. Superparasitism prolonged development time and increased mortality of parasitoid progeny, reduced body size and longevity of the ensuing adults, compared to single parasitism. But such fitness costs of superparasitism can vary with host species, instars, and plants on which the host larvae fed. The development time of parasitoid progeny was shorter and adult size smaller thanin superparasitized hosts of Spodoptera exigua (Hiibner) than Prodenia litura (Fabricius) and Helicoverpa Armigera (Hiibner)(Lepidoptera:Noctuidae). The development time was extendedand adult body size and lifespan increased with increasing host instars at superparasitism, but the progeny survival was higher in middle instars than earlier and later instars. A significant interaction on body size was found between parasitic mode and host plants, where body size of parasitoid progeny was22.5%smaller in superparasitized hosts than in single parasitized larvae when the host larvae fed on cotton, but25.0%when the hosts fed on cabbage. The experiment results suggested that fitness costs of superparasitism to parasitoids can vary with host quality.
3Experiments were carried out to evaluate effects of intraspecific competition, egg load and host encounter rate on the propensity of superparasitism. The analyses, using log-linear model with negative binomial distribution, showed that parasitoids with high host-encounter rate performed more superparasitism than those with low host-encounter rate. An interaction was found between competition and encounter rate with the effect on superparasitism, where parasitoids with high encounter rate performed more superparasitism when they experienced strong competition, but the contrary was true when they had an experience of no-competition. An interaction between competition and egg load was also exhibited. Time of host discrimination was used to evaluate hazard of host being superparasitized by COX proportional hazard model, which showed that host encounter rate and its interaction with egg load had significant impacts on instant probability of superparasitism. The results suggested that potential factors such as intra-specific competition, host abundance, and egg load may often be interacted to influence the propensity of superparasitism.
4Dual-choice trials were performed to observe preferences of M. pulchricornis between healthy and parasitized hosts as affected by parasitization experience and interval time after parasitization. The choice trials showed significant effects of parasitization experience and interval time after parasitization. The parasitoids without an experience of parasitization showed high propensity to attack parasitized hosts than those with the experience of parasitizing healthy hosts. The probability of superparasitism decreased with extending of interval time after parasitization. The analysis of recognition time using COX proportional hazard model indicated diminishing hazards of being superparasitization with extending of interval time after single parasitism. The results suggested that M. pulchricornis have the ability of discrimination between healthy and parasitized hosts, which can obviously be improved by experiences of parasitization.
5A release-recapture method was used to study superparastism as affected by host stages and body weight in the soybean field. The results showed an average parasitism of54.2%(168/310) and superparasitism of23.9%(74/310). The extent of superparasitism, as measured by the number of eggs laid in a host, was negatively influenced by host body weight. The probability of single parasitism decreased with host instars and body weight. But the probability of superparasitism increased with host instars and did not change with host body weight. The experiments suggested that superparasitism could happen in the filed, which may be associated with host life stages at the time of oviposition.
6The recognition time is a prerequisite for evaluation of host quality prior to oviposition in parasitoids. It was measured and analyzed using the COX proportional hazard model for the study of superparasitism hazards (conditional probabilty) as affected by covariate facters. The study suggested that the recognition time used by parasitoids prior to superparasitism can a function of acceptance of the hosts, in that the longer the recognition time the more reluctant the parasitoids to superparasitize the hosts, which suggests a less hazard of being parasitized for the hosts. The current study suggests a noval chacteristic that can be useful for the assessment of superparasitism in parasitoids.
1为探究斑痣悬茧蜂在寄主斑块上的驻留时间与过寄生程度(产卵数)的相关性,采用单选试验设计,观察和分析了斑痣悬茧蜂在斑块内的停留时间与其在寄主幼虫体内产卵数的关系。结果表明,寄生蜂在斑块内驻留的时间影响寄生蜂在每头寄主上的产卵数,1-3h内的产卵平均数分别为0.77、0.9、2.5粒；8h的产卵平均数增加到5.8粒。根据研究推断,在高品质(未被寄生)寄主缺乏的情况下,斑痣悬茧蜂倾向于过寄生。
2为探究寄主品质如何影响过寄生对子代蜂适合度的胁迫作用,采用单选试验设计,研究了以下因素对过寄生行为及其子代发育的影响：不同种寄主(甜菜夜蛾、棉铃虫和斜纹夜蛾),同一种寄主不同龄期寄主幼虫以及取食不同寄主植物。结果表明,过寄生不利于子代蜂的生长发育,导致子代蜂发育历期延长、死亡率增加、体型变小,寿命变短。发育适合度的损耗在不同种类寄主、不同龄期、取食不同植物寄主间发生变化。过寄生甜菜夜蛾幼虫羽化的子代蜂的发育历期、后足胫节比过寄生棉铃虫、斜纹夜羽化的子代蜂的短。与对照不同,过寄生不同龄期的寄主幼虫的子代蜂发育历期、后足胫节、寿命随着龄期的增加而增加；但中间龄期羽化子代蜂的存活率高于从其他龄期羽化的子代蜂。过寄生寄主幼虫取食大豆、棉花和小白菜后的子代蜂发育适合度均显著降低,寄生方式与寄主取食的植物对子代蜂的体型大小有互作效应,与对照相比,从过寄生取食大豆的寄主幼虫中羽化的子代蜂体型减小了22.8%；从过寄生取食棉花的寄主幼虫中羽化的子代蜂体型减小了22.5%；从过寄生取食小白菜的寄主幼虫羽化的子代蜂体型减小了25.0%。据试验研究结果推论,过寄生对寄生蜂适合度的不利影响随寄主品质而变化。
3为探究竞争、寄主充裕程度、卵限制等因素如何影响寄生蜂的过寄生行为,观察了种内竞争、寄主相遇率、寄生蜂抱卵量等对过寄生行为的影响。用基于负二项分布的对数线性模型分析影响过寄生的因素结果表明,种内竞争不影响过寄生发生数量；高抱卵量寄生蜂过寄生寄主的数量高于低抱卵量寄生蜂；寄主相遇率影响过寄生数量,表现为与寄主相遇率高的寄生蜂发生过寄生的次数显著高于与寄主相遇率低的寄生蜂；种内竞争与寄主相遇率对寄生蜂过寄生寄主的数量有互作效应。用识别时间估计寄主被过寄生的风险分析表明,寄主相遇率影响寄主被过寄生的概率,抱卵量和寄主相遇率对过寄生概率具有显著互作影响。据试验结果推论,种内竞争、寄主丰盛度和抱卵量等潜在因素通常互作影响过寄生的发生。
4为评价斑痣悬茧蜂的寄主鉴别能力及其影响因素,采用双选试验研究了寄生蜂的寄生经历和寄主被寄生后的间隔时间对斑痣悬茧蜂在健康寄主和被寄生寄主的选择性。对选择频次进行的分析表明,寄主被寄生后的间隔时间和寄生蜂的寄生经历均对过寄生发生的概率均有显著影响,过寄生概率随寄主被寄生后的间隔时间延长而降低；有寄生经历的寄生蜂发生过寄生的概率低于无寄生经历的寄生蜂。首次测度寄生蜂的寄主识别时间并用COX比例风险模型进行拟合表明,发生过寄生的风险随寄主被寄生后间隔时间的延长而减小,因寄生蜂具有过寄生经历而减小,但寄生蜂具有单寄生经历对过寄生风险没有显著影响。据研究结果推论,斑痣悬茧蜂不仅能够辨别被寄生寄主,而且该能力随寄主被初次寄生后间隔时间延长以及寄生经历而增强。
5野外释放-回收试验的结果表明,斑痣悬茧蜂寄生率为54.2%(168/3l0),过寄生率为23.9%(74/310),寄主体重对寄生蜂在单头寄主上的产卵数有显著影响,随着寄主体重的增加而降低；寄生概率随寄主龄期的增加而降低,但过寄生概率随着寄主龄期的增大而升高；寄生概率随着体重增大而降低,但过寄生概率不受寄主体重影响。据调查结果推论,野外环境中斑痣悬茧蜂可能发生过寄生,而且过寄生程度与产卵时寄主发育时期有关。
6寄生蜂对寄主的识别时间长短反映了其对寄主品质的评价结果。本研究观察测定了斑痣悬茧蜂在评价寄主中所用的识别时间,利用COX比例风险模型拟合识别时间与相关因素的关系,以估计这些因素对过寄生发生倾向的影响。研究结果表明,寄生蜂在决定进行过寄生之前评价寄主所用的识别时间,反映了其对寄主的接受程度,用时长表明寄主质量较差(过寄生),意味着该寄主被寄生的风险较低。研究结果说明,观察寄生蜂在辨别寄主中的识别时间为评价其过寄生行为提供了一个新的特征。
Parasitoid deposits a clutch of eggs (the clutch may consist of a single egg) into a host that has already been parasitized by a member of the same species or itself, a behavior called "superparasitism", and the ensuing larval competition ends in the death of all but one. Superparasitism was not only a focus of theoretical study in evolutionary ecology to understand its adaptation, but also an issue in the practice of biological control using parasitoids to control pests in agriculture. Superparasitism is a common phenomenon in solitary parasitoids, but so far the mechanism of superparasitism is still open to debate. Using Meteorus pulchricornis (Wesmael)(Hymenoptera:Braconidae) as model system, this study was conducted to address four questions:i) whether the parasitoid can discriminate between healthy and parasitized hosts; ii) whether the fitness costs of superparasitism on parasitids vary with host quality; iii) whether potential factors interact to affect superparasitism; iv) whether time-associated behavioral traits might be utilized to evaluate the propensity of superparasitism.The main results and conclusions are summarized as follows.
1A non-choice test was performed to examine the relationship between residence time on host patches and extent of superparasitism as measured by number of eggs per host. The results showed that the residence time had a significant effect on the number of eggs laid in a host. When the residence time was from1to3hours, an average of eggs in a host was0.77,0.9, and2.5respectively, but when the residence time was8hours, an average of5.8eggs was laid in a host. The study suggested that M. pulchricornis have an increasing tendency toward superparasitism with decreasing availability of high-quality hosts.
2In order to examine effects of host quality attributes on fitness consequences of superparasitism for development of parasitoid progeny. A single choice test was used to observe influences of host species, instars, and plant species respectively on the host suitability in superparasitism for the development of parasitoid progeny in a solitary endoparasitoid, M. pulchricornis. Superparasitism prolonged development time and increased mortality of parasitoid progeny, reduced body size and longevity of the ensuing adults, compared to single parasitism. But such fitness costs of superparasitism can vary with host species, instars, and plants on which the host larvae fed. The development time of parasitoid progeny was shorter and adult size smaller thanin superparasitized hosts of Spodoptera exigua (Hiibner) than Prodenia litura (Fabricius) and Helicoverpa Armigera (Hiibner)(Lepidoptera:Noctuidae). The development time was extendedand adult body size and lifespan increased with increasing host instars at superparasitism, but the progeny survival was higher in middle instars than earlier and later instars. A significant interaction on body size was found between parasitic mode and host plants, where body size of parasitoid progeny was22.5%smaller in superparasitized hosts than in single parasitized larvae when the host larvae fed on cotton, but25.0%when the hosts fed on cabbage. The experiment results suggested that fitness costs of superparasitism to parasitoids can vary with host quality.
3Experiments were carried out to evaluate effects of intraspecific competition, egg load and host encounter rate on the propensity of superparasitism. The analyses, using log-linear model with negative binomial distribution, showed that parasitoids with high host-encounter rate performed more superparasitism than those with low host-encounter rate. An interaction was found between competition and encounter rate with the effect on superparasitism, where parasitoids with high encounter rate performed more superparasitism when they experienced strong competition, but the contrary was true when they had an experience of no-competition. An interaction between competition and egg load was also exhibited. Time of host discrimination was used to evaluate hazard of host being superparasitized by COX proportional hazard model, which showed that host encounter rate and its interaction with egg load had significant impacts on instant probability of superparasitism. The results suggested that potential factors such as intra-specific competition, host abundance, and egg load may often be interacted to influence the propensity of superparasitism.
4Dual-choice trials were performed to observe preferences of M. pulchricornis between healthy and parasitized hosts as affected by parasitization experience and interval time after parasitization. The choice trials showed significant effects of parasitization experience and interval time after parasitization. The parasitoids without an experience of parasitization showed high propensity to attack parasitized hosts than those with the experience of parasitizing healthy hosts. The probability of superparasitism decreased with extending of interval time after parasitization. The analysis of recognition time using COX proportional hazard model indicated diminishing hazards of being superparasitization with extending of interval time after single parasitism. The results suggested that M. pulchricornis have the ability of discrimination between healthy and parasitized hosts, which can obviously be improved by experiences of parasitization.
5A release-recapture method was used to study superparastism as affected by host stages and body weight in the soybean field. The results showed an average parasitism of54.2%(168/310) and superparasitism of23.9%(74/310). The extent of superparasitism, as measured by the number of eggs laid in a host, was negatively influenced by host body weight. The probability of single parasitism decreased with host instars and body weight. But the probability of superparasitism increased with host instars and did not change with host body weight. The experiments suggested that superparasitism could happen in the filed, which may be associated with host life stages at the time of oviposition.
6The recognition time is a prerequisite for evaluation of host quality prior to oviposition in parasitoids. It was measured and analyzed using the COX proportional hazard model for the study of superparasitism hazards (conditional probabilty) as affected by covariate facters. The study suggested that the recognition time used by parasitoids prior to superparasitism can a function of acceptance of the hosts, in that the longer the recognition time the more reluctant the parasitoids to superparasitize the hosts, which suggests a less hazard of being parasitized for the hosts. The current study suggests a noval chacteristic that can be useful for the assessment of superparasitism in parasitoids.
引文
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