本文選題：白蠟吉丁腫腿蜂　切入點：寄主適應(yīng)　出處：《中國林業(yè)科學(xué)研究院》2016年博士論文　論文類型：學(xué)位論文

【摘要】：白蠟吉丁腫腿蜂Sclerodermus pupariae Yang et Yao(Hymenoptera:Bethylidae)最早被發(fā)現(xiàn)自然寄生白蠟窄吉丁Agrilus planipennis(Coleoptera:Buprestidae)幼蟲和蛹。后被證實可寄生多種吉丁甲科和天�？评ハx的幼蟲,是一種多寄主型抑性外寄生蜂。該寄生蜂具有搜索能力強、發(fā)育周期短、繁殖潛力大和種群極度偏雌等特點。為了研究該寄生蜂應(yīng)對環(huán)境變化的適應(yīng)能力,進而優(yōu)化其大規(guī)模繁殖技術(shù)和高效的利用于防治不同種類的鉆蛀性害蟲。本文研究了當(dāng)寄主條件和生活環(huán)境變化時白蠟吉丁腫腿蜂的行為和生理變化規(guī)律,揭示了白蠟吉丁腫腿蜂應(yīng)對復(fù)雜多變的環(huán)境的生存能力和適應(yīng)機制,證明了其是一種優(yōu)秀天敵,可被用于多種林業(yè)蛀干害蟲生物防治。主要研究結(jié)果如下:1.白蠟吉丁腫腿蜂因寄生寄主不同可形成對某一種寄主的專性適應(yīng)。長期自然寄生于白蠟窄吉丁幼蟲的寄生蜂同時面臨其自然寄主白蠟窄吉丁和非自然寄主栗山天牛Massicus raddei(Coleoptera:Cerambycidae)時表現(xiàn)為對前者有極強的選擇和寄生行為。然而,隨著寄生蜂在栗山天牛上繁殖的世代數(shù)增加,腫腿蜂雌蜂開始表現(xiàn)出對栗山天牛幼蟲有逐漸增強的選擇寄生行為。連續(xù)以栗山天牛為寄主繁殖至第11代的寄生蜂和自然寄生原始寄主白蠟窄吉丁的寄生蜂雌蜂在無選擇試驗中對栗山天牛和白蠟窄吉丁兩種寄主適應(yīng)性差異顯著。長期寄生栗山天牛的寄生蜂繼續(xù)以栗山天牛作寄主的適合度更高,而長期寄生白蠟窄吉丁的寄生蜂以原始寄主繁殖的適合度更高。同樣,兩個群體的寄生蜂對非自然寄主栗山天牛幼蟲搜索和寄生能力也差異顯著。長期以栗山天牛作寄主的寄生蜂可更高效的發(fā)現(xiàn)和成功寄生寄主植物內(nèi)部的栗山天牛幼蟲。2.寄生蜂成蟲期的學(xué)習(xí)經(jīng)歷可以使它們很快的適應(yīng)一種新的環(huán)境。寄生蜂成蟲早期經(jīng)歷可以增強寄生蜂對相關(guān)環(huán)境的辨識能力。成蟲早期經(jīng)歷主要來源于寄生蜂羽化時通過啃食繭殼而獲得的寄主遺留在寄生蜂繭殼上的化學(xué)信息物質(zhì)。寄生蜂成蟲后期的寄主相關(guān)學(xué)習(xí)經(jīng)歷對其寄主選擇行為影響十分強烈。在成蟲期優(yōu)先接觸的學(xué)習(xí)環(huán)境會促使它們隨后偏好選擇前期所經(jīng)歷過的相關(guān)寄主。并且,成蟲期的取食經(jīng)歷對腫腿蜂的寄主選擇行為影響很大,取食某一種寄主后,在隨后氣味源選擇中對該寄主偏好性更強。成蟲期的學(xué)習(xí)經(jīng)歷還會顯著提高寄生蜂對前期經(jīng)歷過的寄主的寄生能力。3.白蠟吉丁腫腿蜂應(yīng)對不同大小寄主可權(quán)衡生殖和發(fā)育過程。寄生蜂的產(chǎn)卵量主要由寄主條件和產(chǎn)卵前寄生蜂自身的生理狀態(tài)決定,腫腿蜂寄生中等個體大小的栗山天牛低齡幼蟲時產(chǎn)卵量最大。由于寄主質(zhì)量的不均等,母蜂產(chǎn)出不同數(shù)量的卵在相應(yīng)寄主上而引起了子代蜂發(fā)育規(guī)律的多態(tài)性。在小寄主上繁殖的幼期寄生蜂所能獲得的營養(yǎng)有限,所以可在更短的時間內(nèi)能發(fā)育至成熟,但是最終卻只能獲得相對較小的個體質(zhì)量;相反,在個體更大的寄主上繁殖的寄生蜂可獲得相對更大的個體質(zhì)量,但會顯著的延長其幼期周期。寄生蜂依據(jù)寄主營養(yǎng)條件的變化,協(xié)調(diào)自身發(fā)育進程,以此來平衡發(fā)育周期和個體大小兩者之間的矛盾。4.雌蜂密度增大提高了對寄主開發(fā)利用的成功率。當(dāng)一頭寄主上接入的母代腫腿蜂數(shù)量增多時,寄主被成功麻痹制服所需時間大大縮短,并且寄生蜂開始產(chǎn)卵前所需的時間也顯著縮短。母代寄生蜂數(shù)量增大,減少了因為寄主抵御行為而引起的寄生蜂死亡,因此寄生蜂群體的產(chǎn)卵成功率提高。當(dāng)雌蜂數(shù)量由1頭增加至4頭時,在單頭寄主上寄生蜂總的產(chǎn)卵量表現(xiàn)為逐步增加,并且子代數(shù)量也由94頭增加至164頭。子代數(shù)量的增多表明了白蠟吉丁腫腿蜂個體間同類相食的情況很少出現(xiàn)。雖然當(dāng)子代數(shù)量增加時,由于寄主資源的限定而是得單頭寄生蜂的個體大小有輕微降低。但是從寄生蜂群體局部資源增加理論來說,這種多頭母蜂開發(fā)一頭寄主的情況時有利的。5.應(yīng)對寄主資源競爭,腫腿蜂可發(fā)育為更多的有翅型個體而具備隨時遷出不利環(huán)境的能力。隨著在一頭寄主上繁殖的幼期子代蜂數(shù)量增多,其最終成蜂群體里面的有翅型個體也相應(yīng)增多,表明了營養(yǎng)競爭是促進腫腿蜂產(chǎn)生有翅型后代的重要因素。有翅型或無翅型白蠟吉丁腫腿蜂對寄主的寄生能力無顯著差異,表明了在幼期發(fā)育過程中更多的能量分配給翅的發(fā)育并未對具翅型個體生命活力造成負面的效應(yīng)。未發(fā)現(xiàn)有翅型蜂在個體大小和壽命上較無翅個體有顯著的差異。腫腿蜂子代雌性不是簡單地遵循親本的翅的表現(xiàn)形式,而且無翅型雌蜂略偏向產(chǎn)出更多的有翅型雌性后代。6.白蠟吉丁腫腿蜂可以調(diào)整自身寄主結(jié)構(gòu)而應(yīng)對寄主資源外競爭的關(guān)系。對白蠟吉丁腫腿蜂而言,其可利用整個幼期的光肩星天牛anoplophoraglabripennis(coleoptera:cerambycidae)(低齡幼蟲、老熟幼蟲和蛹)作為其生殖產(chǎn)卵的寄主,而另外一種跟它競爭寄主的天敵花絨寄甲主要寄生天牛老熟幼蟲和蛹。腫腿蜂和寄甲成蟲同時存在,腫腿蜂最終可利用的寄主數(shù)量比花絨寄甲更多,前者對寄主寄生率較后者高10%。若腫腿蜂與寄甲卵被同時使用,寄甲卵可迅速孵化為幼蟲而開始寄主搜索過程,因此最終被花絨寄甲寄生的寄主較腫腿蜂更多,此時寄甲對寄主的寄生率較腫腿蜂高13%左右。但總的來說,白蠟吉丁腫腿蜂并未出現(xiàn)由于花絨寄甲與其競爭寄主而出現(xiàn)不能成功延續(xù)種群繁殖。此外,未發(fā)現(xiàn)寄主有被兩種天敵重寄生的現(xiàn)象,表明兩種具備同時被釋放于林間用于生物防治的生態(tài)基礎(chǔ)。7.非寄主類糖類營養(yǎng)有助于腫腿蜂應(yīng)對林間寄主缺乏的情況。糖類物質(zhì)可以作為供腫腿蜂補充營養(yǎng)的潛在食物源。寄主血淋巴是作為寄生蜂成蟲的固有的營養(yǎng)資源,但發(fā)現(xiàn)寄生蜂在僅補充蜂蜜的情況下也可以獲得對寄主寄生能力的增強和自身壽命的延長。剛羽化的腫腿蜂在獲得取食蜂蜜經(jīng)歷后,其單次產(chǎn)卵量均較無補充營養(yǎng)經(jīng)歷的高20%,并且其壽命顯著延長。同時,試驗結(jié)果證明了取食蜂蜜或寄主血淋巴對腫腿蜂生殖和壽命的增強作用相近。在林間如果遇到某一段時期寄主資源相對缺乏的時候,腫腿蜂可利用這些營養(yǎng)物質(zhì)延長壽命,保存在林間的種群數(shù)量以此實現(xiàn)后期對出現(xiàn)合適寄主時的成功開發(fā)。8.白蠟吉丁腫腿蜂具有撫幼行為習(xí)性,可以協(xié)助幼蜂更好的完成發(fā)育。當(dāng)發(fā)育在寄主上的白蠟吉丁腫腿蜂幼蜂數(shù)量過多,此種情況下幼蟲化蛹前無母蜂協(xié)助便會出現(xiàn)大量的死亡。死亡的主要原因為一定數(shù)量的老熟幼蟲因自身不能移動而堆積,因此不能成功化蛹。但是若環(huán)境中有雌蜂存在,雌蜂會協(xié)助幼蜂移動,幼期死亡率會大大降低。親本撫育行為的重要時期體現(xiàn)在腫腿蜂幼蟲期,而其余時期即使無母蜂陪伴寄生蜂發(fā)育也不會受較大影響。腫腿蜂的撫幼行為同樣還存在于母蜂對非自產(chǎn)幼蜂的撫育,并且這種撫幼行為不會受到是否具有產(chǎn)卵經(jīng)歷的影響。本文深入研究了多寄主型寄生蜂白蠟吉丁腫腿蜂依據(jù)其固有的行為和發(fā)育可塑性而具備快速適應(yīng)變化多端的環(huán)境的能力。首先,白蠟吉丁腫腿蜂可通過寄生不用寄主而獲得對某一寄主的專性適應(yīng)能力,并且通過短期的寄主相關(guān)學(xué)習(xí)經(jīng)歷也同樣可以快速的適應(yīng)變化的寄主和寄主棲境環(huán)境。同時還探明了白蠟吉丁腫腿蜂在應(yīng)對寄主資源缺乏時的適應(yīng)機制。多頭寄生蜂同時寄生一頭寄主會造成種內(nèi)資源競爭,但同時對寄主資源的開發(fā)更有利,并且腫腿蜂在應(yīng)對和花絨寄甲的種間資源競爭時可調(diào)整自身寄主結(jié)構(gòu)而確保種群的延續(xù)。寄生蜂面對寄主資源缺乏時可通過取食花蜜獲得壽命的延長,同時棲境內(nèi)密度過大可誘導(dǎo)更多寄生蜂發(fā)育為有翅型個體而獲得可隨時遷出不利環(huán)境的能力。最后,白蠟吉丁腫腿蜂的撫幼行為習(xí)性確保了幼期寄生蜂在不利環(huán)境中的發(fā)育成功率。我們的研究結(jié)果表明了白蠟吉丁腫腿蜂有著極強的行為和生理可塑性,它們具備應(yīng)對復(fù)雜多變環(huán)境的快速適應(yīng)能力,是一種可被深入開發(fā)和應(yīng)用于多種天牛類和吉丁甲類昆蟲的生物防治的優(yōu)勢天敵。
[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.

【學(xué)位授予單位】：中國林業(yè)科學(xué)研究院
【學(xué)位級別】：博士
【學(xué)位授予年份】：2016
【分類號】：S476.3
，

本文編號：1632389