多寄主型寄生蜂白蜡吉丁肿腿蜂应对环境变化的行为和发育策略研究

发布时间：2018-03-19 02:19

  本文选题：白蜡吉丁肿腿蜂　切入点：寄主适应　出处：《中国林业科学研究院》2016年博士论文　论文类型：学位论文

【摘要】：白蜡吉丁肿腿蜂Sclerodermus pupariae Yang et Yao(Hymenoptera:Bethylidae)最早被发现自然寄生白蜡窄吉丁Agrilus planipennis(Coleoptera:Buprestidae)幼虫和蛹。后被证实可寄生多种吉丁甲科和天牛科昆虫的幼虫,是一种多寄主型抑性外寄生蜂。该寄生蜂具有搜索能力强、发育周期短、繁殖潜力大和种群极度偏雌等特点。为了研究该寄生蜂应对环境变化的适应能力,进而优化其大规模繁殖技术和高效的利用于防治不同种类的钻蛀性害虫。本文研究了当寄主条件和生活环境变化时白蜡吉丁肿腿蜂的行为和生理变化规律,揭示了白蜡吉丁肿腿蜂应对复杂多变的环境的生存能力和适应机制,证明了其是一种优秀天敌,可被用于多种林业蛀干害虫生物防治。主要研究结果如下:1.白蜡吉丁肿腿蜂因寄生寄主不同可形成对某一种寄主的专性适应。长期自然寄生于白蜡窄吉丁幼虫的寄生蜂同时面临其自然寄主白蜡窄吉丁和非自然寄主栗山天牛Massicus raddei(Coleoptera:Cerambycidae)时表现为对前者有极强的选择和寄生行为。然而,随着寄生蜂在栗山天牛上繁殖的世代数增加,肿腿蜂雌蜂开始表现出对栗山天牛幼虫有逐渐增强的选择寄生行为。连续以栗山天牛为寄主繁殖至第11代的寄生蜂和自然寄生原始寄主白蜡窄吉丁的寄生蜂雌蜂在无选择试验中对栗山天牛和白蜡窄吉丁两种寄主适应性差异显著。长期寄生栗山天牛的寄生蜂继续以栗山天牛作寄主的适合度更高,而长期寄生白蜡窄吉丁的寄生蜂以原始寄主繁殖的适合度更高。同样,两个群体的寄生蜂对非自然寄主栗山天牛幼虫搜索和寄生能力也差异显著。长期以栗山天牛作寄主的寄生蜂可更高效的发现和成功寄生寄主植物内部的栗山天牛幼虫。2.寄生蜂成虫期的学习经历可以使它们很快的适应一种新的环境。寄生蜂成虫早期经历可以增强寄生蜂对相关环境的辨识能力。成虫早期经历主要来源于寄生蜂羽化时通过啃食茧壳而获得的寄主遗留在寄生蜂茧壳上的化学信息物质。寄生蜂成虫后期的寄主相关学习经历对其寄主选择行为影响十分强烈。在成虫期优先接触的学习环境会促使它们随后偏好选择前期所经历过的相关寄主。并且,成虫期的取食经历对肿腿蜂的寄主选择行为影响很大,取食某一种寄主后,在随后气味源选择中对该寄主偏好性更强。成虫期的学习经历还会显著提高寄生蜂对前期经历过的寄主的寄生能力。3.白蜡吉丁肿腿蜂应对不同大小寄主可权衡生殖和发育过程。寄生蜂的产卵量主要由寄主条件和产卵前寄生蜂自身的生理状态决定,肿腿蜂寄生中等个体大小的栗山天牛低龄幼虫时产卵量最大。由于寄主质量的不均等,母蜂产出不同数量的卵在相应寄主上而引起了子代蜂发育规律的多态性。在小寄主上繁殖的幼期寄生蜂所能获得的营养有限,所以可在更短的时间内能发育至成熟,但是最终却只能获得相对较小的个体质量;相反,在个体更大的寄主上繁殖的寄生蜂可获得相对更大的个体质量,但会显著的延长其幼期周期。寄生蜂依据寄主营养条件的变化,协调自身发育进程,以此来平衡发育周期和个体大小两者之间的矛盾。4.雌蜂密度增大提高了对寄主开发利用的成功率。当一头寄主上接入的母代肿腿蜂数量增多时,寄主被成功麻痹制服所需时间大大缩短,并且寄生蜂开始产卵前所需的时间也显著缩短。母代寄生蜂数量增大,减少了因为寄主抵御行为而引起的寄生蜂死亡,因此寄生蜂群体的产卵成功率提高。当雌蜂数量由1头增加至4头时,在单头寄主上寄生蜂总的产卵量表现为逐步增加,并且子代数量也由94头增加至164头。子代数量的增多表明了白蜡吉丁肿腿蜂个体间同类相食的情况很少出现。虽然当子代数量增加时,由于寄主资源的限定而是得单头寄生蜂的个体大小有轻微降低。但是从寄生蜂群体局部资源增加理论来说,这种多头母蜂开发一头寄主的情况时有利的。5.应对寄主资源竞争,肿腿蜂可发育为更多的有翅型个体而具备随时迁出不利环境的能力。随着在一头寄主上繁殖的幼期子代蜂数量增多,其最终成蜂群体里面的有翅型个体也相应增多,表明了营养竞争是促进肿腿蜂产生有翅型后代的重要因素。有翅型或无翅型白蜡吉丁肿腿蜂对寄主的寄生能力无显著差异,表明了在幼期发育过程中更多的能量分配给翅的发育并未对具翅型个体生命活力造成负面的效应。未发现有翅型蜂在个体大小和寿命上较无翅个体有显著的差异。肿腿蜂子代雌性不是简单地遵循亲本的翅的表现形式,而且无翅型雌蜂略偏向产出更多的有翅型雌性后代。6.白蜡吉丁肿腿蜂可以调整自身寄主结构而应对寄主资源外竞争的关系。对白蜡吉丁肿腿蜂而言,其可利用整个幼期的光肩星天牛anoplophoraglabripennis(coleoptera:cerambycidae)(低龄幼虫、老熟幼虫和蛹)作为其生殖产卵的寄主,而另外一种跟它竞争寄主的天敌花绒寄甲主要寄生天牛老熟幼虫和蛹。肿腿蜂和寄甲成虫同时存在,肿腿蜂最终可利用的寄主数量比花绒寄甲更多,前者对寄主寄生率较后者高10%。若肿腿蜂与寄甲卵被同时使用,寄甲卵可迅速孵化为幼虫而开始寄主搜索过程,因此最终被花绒寄甲寄生的寄主较肿腿蜂更多,此时寄甲对寄主的寄生率较肿腿蜂高13%左右。但总的来说,白蜡吉丁肿腿蜂并未出现由于花绒寄甲与其竞争寄主而出现不能成功延续种群繁殖。此外,未发现寄主有被两种天敌重寄生的现象,表明两种具备同时被释放于林间用于生物防治的生态基础。7.非寄主类糖类营养有助于肿腿蜂应对林间寄主缺乏的情况。糖类物质可以作为供肿腿蜂补充营养的潜在食物源。寄主血淋巴是作为寄生蜂成虫的固有的营养资源,但发现寄生蜂在仅补充蜂蜜的情况下也可以获得对寄主寄生能力的增强和自身寿命的延长。刚羽化的肿腿蜂在获得取食蜂蜜经历后,其单次产卵量均较无补充营养经历的高20%,并且其寿命显著延长。同时,试验结果证明了取食蜂蜜或寄主血淋巴对肿腿蜂生殖和寿命的增强作用相近。在林间如果遇到某一段时期寄主资源相对缺乏的时候,肿腿蜂可利用这些营养物质延长寿命,保存在林间的种群数量以此实现后期对出现合适寄主时的成功开发。8.白蜡吉丁肿腿蜂具有抚幼行为习性,可以协助幼蜂更好的完成发育。当发育在寄主上的白蜡吉丁肿腿蜂幼蜂数量过多,此种情况下幼虫化蛹前无母蜂协助便会出现大量的死亡。死亡的主要原因为一定数量的老熟幼虫因自身不能移动而堆积,因此不能成功化蛹。但是若环境中有雌蜂存在,雌蜂会协助幼蜂移动,幼期死亡率会大大降低。亲本抚育行为的重要时期体现在肿腿蜂幼虫期,而其余时期即使无母蜂陪伴寄生蜂发育也不会受较大影响。肿腿蜂的抚幼行为同样还存在于母蜂对非自产幼蜂的抚育,并且这种抚幼行为不会受到是否具有产卵经历的影响。本文深入研究了多寄主型寄生蜂白蜡吉丁肿腿蜂依据其固有的行为和发育可塑性而具备快速适应变化多端的环境的能力。首先,白蜡吉丁肿腿蜂可通过寄生不用寄主而获得对某一寄主的专性适应能力,并且通过短期的寄主相关学习经历也同样可以快速的适应变化的寄主和寄主栖境环境。同时还探明了白蜡吉丁肿腿蜂在应对寄主资源缺乏时的适应机制。多头寄生蜂同时寄生一头寄主会造成种内资源竞争,但同时对寄主资源的开发更有利,并且肿腿蜂在应对和花绒寄甲的种间资源竞争时可调整自身寄主结构而确保种群的延续。寄生蜂面对寄主资源缺乏时可通过取食花蜜获得寿命的延长,同时栖境内密度过大可诱导更多寄生蜂发育为有翅型个体而获得可随时迁出不利环境的能力。最后,白蜡吉丁肿腿蜂的抚幼行为习性确保了幼期寄生蜂在不利环境中的发育成功率。我们的研究结果表明了白蜡吉丁肿腿蜂有着极强的行为和生理可塑性,它们具备应对复杂多变环境的快速适应能力,是一种可被深入开发和应用于多种天牛类和吉丁甲类昆虫的生物防治的优势天敌。
[Abstract]:The ash borer s.guani Sclerodermus pupariae Yang et Yao (Hymenoptera:Bethylidae) is the earliest discovered natural parasitic Agrilus planipennis Agrilus planipennis (Coleoptera:Buprestidae) larvae and pupae. After being confirmed a variety of parasitic Buprestidae and Cerambycidae larvae, is a type of tumor host parasitoid. The parasitic wasps have search strong ability, short development cycle, reproductive potential and population extremely partial female characteristics. In order to study the parasitoids in response to changes in the environment adaptability, and to optimize its technology and efficient use in prevention and treatment of different types of the borer pests of mass reproduction. This paper studies the behavior when the host conditions and living environment change of ash borer bethylid and changes of physiology, reveals the ash borer s.guani response to the complex environment of the survival ability and adaptation mechanism, it is proved to be a good Show natural enemies, can be used for a variety of forestry pest control stem borer. The main results are as follows: 1. ash borer bethylid parasitized by different forms of a host specific adaptation. The parasitoid long-term parasitic on the natural emerald ash borer larvae also faces its natural host and non natural emerald ash borer host Massicus Massicus raddei (Coleoptera:Cerambycidae) showed strong selection and parasitic behavior of the former. However, as the number of generations of parasitoid breeding in the Massicus on the increase of bethylid parasitoids began to show a gradually increasing parasitic behavior of Massicus continuous to Massicus larvae. As the host of the eleventh generation reproduction of parasitoids and parasitism natural original host parasitoid wasp Agrilus planipennis display of Massicus and Agrilus planipennis two host adaptability difference in no choice test The parasitic wasps parasitic. Long term massicusraddei continue to Massicus as a host for a higher degree, while parasitoids long-term. Agrilus planipennis in the original host reproduction for a higher degree. Similarly, the parasitoid of the two groups of non natural host Massicus larvae search and parasitic ability the difference was significant. The Massicus as the host of the parasitic wasp can be more efficient and successful parasitism of host plants were found inside the Massicus parasitoid larvae of.2. in adult learning experience so that they can quickly adapt to a new environment. Parasitoid adults can enhance the early experience of parasitoids on environment related identification the chemical substances information ability. Adults early experience while the main sources obtained from the parasitoid eclosion by eating the cocoon shell left in the host parasitoid cocoons. Host parasitoid adults later learning experience to send The main impact is very strong. In the host adult stage preferential access to learning environment will lead to their subsequent preference early experience. And experience for feeding adult stage s.guani host selection behavior of great influence, feeding a host, in the following selection of the host odor source the preference is stronger. Adult learning experience will significantly improve the early experience of parasitoid host larvae of.3. ash borer s.guani with different size of the host can weigh the reproductive and developmental process. The physiological state of oviposition of parasitic wasps mainly by the host before spawning conditions and parasitoids decision s.guani parasitic medium size Massicus larvae when spawning. Due to host quality not to be equal, the female produce different quantities of eggs in the corresponding host caused development of offspring parasitoids Polymorphism law. Reproduction in small host parasitoid larvae can obtain nutrition is limited, so in a shorter period of time to mature and develop, but eventually only individual quality is relatively small; on the contrary, the parasitoid breeding in the individual more hosts can get bigger the quality of the individual, but they significantly prolong the young cycle. On the basis of host parasitoid changes the nutritional conditions, in order to coordinate their own development process, the contradictions of the.4. density increases the balance between growth period and individual size both increase on host development and success rate. When the number of generation of s.guani a host access increased when the host was successfully required uniform paralysis time is greatly shortened, and the parasitic wasps required to get started before the spawning time is significantly shortened. The increase of the number of parent parasitoids, reduced because the host resisting behavior arising The parasitoid death, so the parasitoid population spawning success rate increased. When the number increased by 1 to 4, the fecundity of parasitic wasps tend to gradually increase in the single host, and the number of offspring from 94 increased to 164 head. More number of offspring that ash Gideon s.guani among individuals cannibalism is rarely the case. Although when the generation numbers increase, due to limited host resources but single head parasitoid individual size slightly decreased. But the increase from the parasitoid group local resource theory,.5. deal with the host resource competition advantage this long development a female host when the bethylid can develop into more winged individuals have always moved out of ability disadvantageous environment. As the number of breeding in a host of young stage of offspring parasitoids increased, the final group inside the winged bee The individual is increasing correspondingly, that is to promote the nutrition competition s.guani important factors of winged offspring. Winged or parasitic ability of fin type bethylid on ash borer host shows no significant difference in larval development in energy allocation to more development and not on the winged wings individual life has the negative effect. Not found in body size and wing type of bee life than wingless individuals have significant difference. Forms of s.guani offspring of female is not simply follow the parent wing, and wingless type females slightly produce more winged female offspring of.6. wax Gideon s.guani can adjust its host structure and deal with host resources outside the competition. The ash borer bethylid, which can use the nepionic anoplophoraglabripennis anoplophoraglabripennis (coleoptera:cerambycidae) (younger The larvae, larvae and pupae) as the fecundity of host, and the other one with its host competition predator Dastarcus helophoroides main parasitic beetles old larvae and pupae. Bethylid and sent a number of adults exist at the same time, the host s.guani eventually available more than Dastarcus helophoroides. The former host of parasitic rate higher than 10%. if bethylid and send a eggs to be used at the same time, send a eggs can be quickly hatched into larvae began to host the search process, thus eventually Dastarcus helophoroides parasitic host than s.guani more, then send a host of parasitic rate of s.guani high about 13%. But in general, the ash borer s.guani did not appear because of Dastarcus helophoroides and its host not successful continuation of population reproduction. In addition, there are two kinds of host not found natural hyperparasitism phenomenon, show that the two have also released in the woods for biological control The ecological basis of.7. treatment of non host carbohydrate class helps s.guani to deal with the lack of sugar. The host material can be used as a nutritional supplement for s.guani potential food source. Host hemolymph as nutrient resources inherent in adult parasitoids, but found that prolonged parasitoids in the case only added honey can also be enhanced to host the parasitic ability and their life. Newly emerged s.guani in honey feeding after the single fecundity were 20% higher nutrition experience, and the life was prolonged. At the same time, the test results proved that honey feeding or host hemolymph of s.guani reproduction and life enhancing effect in the woods when similar. If the host resources of a period of relative lack of s.guani can use these nutrients to prolong life, save in the population of the forest in order to achieve A late nursery behavior of the successful development of suitable host.8. ash borer bethylid, can help young bees better finish development. When the number of ash borer s.guani larvae development in host too much, there will be a large number of deaths in this case before the queen bee larvae pupate and assist. The accumulation of the main causes of death for a certain number of mature larvae due to their own can not move, and therefore can not be successful. But if the pupation environment has the existence, the female will help young bees move, instar'smortality will be greatly reduced. In s.guani larval stage of an important period of parental care behavior, while the remaining period without the queen bee with the parasitoid development is not affected by the behavior of s.guani. Nursery also exists in the young females of non self tending, and this behavior is not affected by whether the nursery has oviposition experience Influence. This in-depth study of the generalist parasitoids of s.guani on the basis of the ash borer inherent behavior and developmental plasticity and have the ability to adapt the environment. Firstly, be the most changeful, bethylid by ash borer was not the host parasitic on a host specific ability to adapt, and by host related learning the short-term experience also can quickly adapt to the change of the host and host habitat environment. At the same time also proved the adaptation mechanism of ash borer s.guani in response to the lack of host resources. At the same time a head of long parasitic wasps parasitic host will cause intraspecific competition for resources, but also the development of host resources better, and swollen Guani send a competition for resources in coping and Dastarcus can adjust its structure and ensure the continuation of the host population. In the face of the host parasitoid lack of resources could be used for nectar feeding Extended life, at the same time the territory of habitat density is too large can induce more parasitoids develop into winged individuals can be obtained at any time from the ability of the disadvantageous environment. Finally, the behaviour of ash nursery s.guani ensure borer development in adverse environments the larval parasitoids success rate. Our results show that the ash borer s.guani has a strong behavioral and physiological plasticity, they have to deal with the complex environment of rapid adaptation, is a day of advantage can be deeply developed and applied in various classes and the insect borer beetle biocontrol of the enemy.

【学位授予单位】：中国林业科学研究院
【学位级别】：博士
【学位授予年份】：2016
【分类号】：S476.3
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本文编号：1632389