寄生蜂寻找隐蔽性寄主害虫的行为机制

林木蛀干类害虫具有高度的隐蔽性,是林业上的一类重要害虫,也是目前世界上最难防治的害虫类群之一.寄生蜂在与寄主长期的协同进化过程中,形成了搜索、发现和攻击寄主害虫的独特机制,能够有效地找到并寄生它们.总结了寄生性天敌寻找、发现并成功定位隐蔽性寄主害虫的行为学机制.寄生蜂可以利用来自嗅觉的化学信息物质(如寄主、寄主粪便、虫道共生菌的挥发性气味)、寄主成虫的化学通讯物质、来自视觉的植物表面色差信息、来自触觉的寄主保护物性状特征、来自寄主取食和运动所产生的介质振动信号以及来自寄主活动和代谢的红外辐射等多种途径有效地发现隐蔽性害虫的位置,从而完成寄生行为.有些寄生蜂还能综合利用来源不同的多种信息,从而提高寄主定位的可靠性和准确性.本文还对寄生蜂寻找寄主的这些线索在生物防治上可能的利用途径和前景进行了讨论.这对促进我国在该领域的研究,充分利用天敌昆虫,提高生物防治效率具有参考价值.     

寄生性天敌寄生行为及研究中应注意的若干问题

作为调节和利用寄生性天敌以控制害虫为害所需的重要基础信息之一,寄生性天敌的寄生行为已有70多年的研究历史。至今在这一领域的很多研究方面。包括寄生性天敌的寄主选择行为、寄主块离开行为、窝卵数以及行为的进化等都取得了很大的进展。本文主要对寄生性天敌的寄主选择行为及寄主块离开行为的最新研究成果进行综述。并就寄生性天敌寄生行为研究过程中经常遇到的一些问题进行讨论。     

多寄主型寄生性天敌昆虫的寄主适应性及其影响因素

寄生性天敌昆虫对不同寄主资源的适应能力是其存活和繁殖的必要条件,它们的寄主选择行为则是其重要的适应结果。多寄主型寄生性天敌昆虫虽然对某种特定寄主资源的利用效率可能不如单寄主型寄生蜂,但却有利于拓展更宽的寄主范围,因此对环境的适应能力更强,更容易在自然界维持其种群的生存。统计结果也表明多寄主型天敌的生物防治效果往往比专食性天敌更高。有时生物防治成功的关键可能并不在于所利用的天敌种类的不同,而在于天敌的不同生物型或地理宗。总结了多寄主型寄生性天敌昆虫的寄主适应能力及其影响因素。寄生性天敌昆虫不仅因地理隔离产生种群分化,也可能因寄生不同的寄主产生种群内的分化,从而更加适应寄主的生活特性和栖境条件。寄主种类、寄主发育阶段、寄主大小、寄主营养、寄主免疫反应、寄主逃避反应、其它天敌的竞争、寄主共生或共栖生物的存在、寄主植物、天敌自身的学习能力及其共生微生物等多种因素对寄生性天敌昆虫的寄主适应性可产生影响。展望了多寄主型寄生性天敌昆虫对新寄主资源的拓展利用能力和适应性在生产上的可能应用前景和途径,以期为明确天敌与寄主间的互作关系,人工驯化寄生性天敌昆虫增强对靶标害虫的控制作用,合理利用天敌提高生物防治效率提供新的思路和理论支持。     

肿腿蜂类寄生蜂室内控害效能评价——以松脊吉丁肿腿蜂为例

为了开发利用天敌资源防治害虫,科学而客观地评价肿腿蜂对蛀干害虫的控害作用,研究了松脊吉丁肿腿蜂(Sclerodermus sp.)对松褐天牛3龄幼虫的功能反应,建立了肿腿蜂类寄生蜂室内控害效能评价体系,并证明了其可行性。该评价体系包括供试虫源标准、寄生蜂生物学与寄生行为学观察、寄生蜂对寄主的功能反应和寻找寄主效应以及寄生蜂的忠岐指数(Y)4个部分。研究结果表明:松脊吉丁肿腿蜂具有较强的控害潜力,单头雌成虫能防治3—4头松褐天牛3龄幼虫,21d内最多可致死9.07头寄主幼虫;寄主密度对单头寄生蜂寄生作用功能反应的测定结果与对寄生作用寻找效应的测定结果以及对产卵量影响的测定结果一致;综合评价该蜂对松褐天牛3龄幼虫控制效能,释放比为1∶1(即1头肿腿蜂雌蜂:1头3龄松褐天牛幼虫)时最高,其忠岐指数Y为39.63,这可能是肿腿蜂类天敌特有的抚幼习性所致,结论也与寄生蜂生物学特性及寄生行为学观察结果相同。研究结论证明该评价体系是稳定可靠的。评价体系在利用天敌昆虫生物防治害虫时为筛选最佳寄生性天敌种类提供了可行的方法和依据。     

补充非寄主源食物显著提高万氏潜蝇姬小蜂孤雌产雌品系对美洲斑潜蝇的控害潜力

[目的]万氏潜蝇姬小蜂Diglyphus wani是潜叶蝇类害虫的重要寄生蜂,可以孤雌产雌生殖,仅产生雌性后代,具有潜在的生防应用价值.该寄生蜂为寄主取食型寄生蜂,自然条件下不仅可取食非寄主源食物,也可直接取食寄主幼虫.本研究旨在探究补充非寄主源食物对其生防效果的影响,从而有效指导其大规模饲养和田间应用.[方法]以万氏...     

六盘山翠金小蜂生物学特性的研究

六盘山翠金小蜂Tritneptis sp.是华北落叶松人工林主要食叶害虫落叶松红腹叶蜂Pristiphora erichsonii Hartig茧期的体外复寄生性天敌。对该蜂的生物学特性研究表明,该蜂在宁夏六盘山1年2代,以幼虫在寄主茧内越冬,翌年4月中旬开始活动取食,5月中旬开始化蛹,成蜂从6月中旬开始羽化,每头虫茧平均出蜂6.14头,雌雄比1.79∶1,雌蜂6月下旬开始产卵,一只雌蜂一生可在12.2头寄主虫茧上产卵,第1代寄生蜂幼虫7月上旬开始孵化并吸食寄主营养发育,7月下旬开始化蛹,8月上旬开始羽化并交尾、产卵寄生。该蜂体型较大,对寄主跟随性强,无重寄生天敌,自然寄生率7.3%～33.6%,是一种优良天敌寄生蜂。     

寄生蜂感受器的结构和功能研究进展

寄生蜂是寄生在寄主体内或体外的一类重要天敌昆虫,在农林害虫生物防治中具有重要作用.寄生蜂对寄主的确定主要依靠触角敲打和产卵器刺探共同完成,作为寄生蜂的重要感觉器官,触角和产卵器上着生有不同种类的感受器,在其寄主搜寻和产卵识别等过程中起着感受外界信息的关键作用.本文综述了国内外有关寄生蜂触角和产卵器上感受器的研究进展,包括感受器的类群划分、超微结构和生理功能等方面,并对该领域中尚待解决的问题和研究展望进行了探讨,以期为深入研究寄生蜂的寄生行为,揭示其对外界信息的感受机制以及将来利用寄生蜂进行害虫的生物防治提供理论基础.     

中国寄生蜂研究及其在害虫生物防治中的应用

寄生蜂是一类重要的寄生性天敌昆虫,种类繁多、习性复杂,在害虫生物防治和综合治理中发挥着极其重要的作用。在产卵时,寄生蜂携带的毒液、多DNA病毒等寄生因子就会随之进入寄主体内,发挥调控寄主生长、发育、免疫、代谢、行为的作用,从而保障了寄生蜂后代的发育。本文主要针对我国寄生蜂的系统分类、资源普查、生物学、生态学、寄主调控、人工繁殖、释放应用、田间保护和助增等方面的基础研究和应用进行了概述和整理。     

红富士苹果园害虫与天敌群落的定量分析

应用多元分析方法对烟台市牟平区红富士苹果园害虫与天敌群落进行了定量分析,采用最优分割法将苹果害虫与天敌群落的时序结构分为5个阶段,并分析了各个时序段害虫及天敌的发生特点.对害虫亚群落、捕食性天敌亚群落和寄生性天敌亚群落进行了主成分和因子分析,明确了不同时期起主要作用的害虫及其天敌种类.典型相关分析表明,主要害虫及其天敌类群之间存在显著相关性,其中金纹细蛾与其寄生蜂、绣线菊蚜与其寄生蜂、山楂叶螨与其专一性捕食性天敌深点颏瓢虫和东方钝绥螨之间的相关性高.     

取食对寄生蜂的影响

寄生蜂是调节昆虫种群动态的重要因子之一,其对寄主害虫产生影响的因素主要有寄生蜂摄入的营养、寄生蜂的寿命和生殖力以及其对寄主的搜寻效率等.寄生蜂的营养主要来源于幼蜂体内营养积累和成蜂取食两个途径.其中成蜂取食又包括取食寄主和取食非寄主食物两个方面.某些种类的成蜂通过取食寄主来获得生存和生殖所需的营养,但是大部分成蜂需要取食糖类来作为其主要能源,而且糖类对寄生蜂的生存和生殖也会产生一定的影响.本文综述了寄生蜂的食物来源、食物营养在寄生蜂体内的分配、取食寄主和非寄主食物对寄生蜂的影响以及寄生蜂在寄主搜寻和非寄主食物搜寻之间的权衡,以期为寄生蜂的饲养和利用提供参考.     

Taxonomy of parasitoid wasps in China: An overview

The parasitoid Hymenoptera are one of the most important groups of natural enemies of agriculture and forestry pests and have been used as biological control agents in IPM programs for a long time. The systematic research on parasitoids and their associations with hosts lays a solid base for biological control of pests. Here we summarize the taxonomic studies undertaken on the parasitoid wasps in China in the last eight decades. At present about 48 families in 12 superfamilies of parasitoid Hymenoptera are known in China, of which the majority of the families have been studied to some extent while a number of them are still poorly studied. Around 6000 species of 32 families which dealt with in this paper are known in China, and some of them have been successfully and widely used in the existing biological control while others are potentially useful in future biocontrol program. This overview provides the international audience a general idea about the current state of the systematic study of parasitoid Hymenoptera conducted in China, the known species of parasitoids and the native and introduced species used in biocontrol in China.     

拟寄生蜂搜索产卵过程中对寄主的竞争

综述拟寄生蜂搜索产卵过程中对寄主竞争的最新研究进展.这类竞争具有四种方式,即标记寄主、杀卵和杀幼、守护寄主和捕食寄主.（1）标记寄主常涉及寄主标记信息素,这是由雌蜂在产卵前、产卵时或产卵后分泌的化学物质.寄主标记信息素常介导拟寄生蜂对已寄生和健康寄主的辨别、减少过寄生和多寄生、减轻种内和种间竞争压力.（2）雌蜂遇到已寄生寄主时,很多种类杀死前一雌蜂遗留的卵和幼虫,再产下自己的卵.雌蜂使用三种方法杀卵和杀幼,即产卵器穿刺、取食和使用有毒物质.通过杀卵和杀幼,产卵雌蜂清除了前一雌蜂遗留的后代,主动改善了寄主品质,从而有利于自身后代的生存.（3）守护寄主在肿腿蜂科、缘腹细蜂科、金小蜂科、缨小蜂科和茧蜂科中均有报道,守护者驱逐入侵者以保护后代及健康寄主.（4）捕食寄主不仅减少了健康寄主数量,且直接导致已寄生寄主中拟寄生蜂卵和幼虫的死亡.雌蜂一般在体内成熟卵量较少时捕食寄主.讨论了研究拟寄生蜂搜索产卵过程中竞争寄主的理论意义和实际应用价值.     

Effect of the primary host for production of both sexes on the mating interaction in an autoparasitoid species

Encarsia sophia (Girault and Dodd) is an autoparasitoid in the hymenopteran family Aphelinidae. The females develop as primary parasitoids on whitefly nymphs (primary hosts), whereas the males develop as hyperparasitoids on their own species or on other primary parasitoid species (secondary hosts). The autoparasitoids not only parasitise whiteflies but also kill them with strong host-feeding capacity. In this study, female and male E. sophia were reared on the primary hosts Trialeurodes vaporariorum and Bemisia tabaci ‘Q’, and the host-feeding and parasitism of wasps on both whitefly species were determined for the four possible different mating combinations: (i) E. sophia females reared on B. tabaci (ESF-BT) mated with E. sophia males from B. tabaci (ESM-BT), (ii) E. sophia females reared on T. vaporariorum (ESF-TV) mated with E. sophia males from T. vaporariorum (ESM-TV), (iii) ESF-BT mated with ESM-TV, and (iv) ESF-TV mated with ESM-BT. ESF-TV mated with ESM-TV killed the largest percentage of whitefly nymphs through host feeding. The ESF-TV with larger body size mating with larger ESM-TV killed more whitefly nymphs through host feeding than those mating with smaller ESM-BT. Whether B. tabaci or T. vaporariorum were used as hosts, ESF-TV mated with ESM-TV and ESM-BT and ESF-BT mated with ESM-BT significantly parasitised more whitefly nymphs than ESF-BT mated with ESM-TV. In general, ESF-BT mated with ESM-TV killed significantly fewer whitefly nymphs through parasitism and host feeding than the other three mating combinations on both whitefly species. These results indicated that the performance of autoparasitoids on insect pests was not only dependent on females but was also affected by mating with males from different primary host species.     

Behavioral mechanisms of parasitic wasps for searching concealed insect hosts

Wood borers are important forest insect pests and difficult to be controlled owing to their concealed behavior. However, parasitic wasps can effectively ascertain and parasitize wood borers as well as other concealed pests by using special searching, finding and attacking mechanisms, which have been developed during the course of long-term coevolution with their hosts. The present paper summarizes the behavioral mechanisms of parasitic wasps involved in searching and locating their concealed hosts. Parasitic wasps can accurately find the location of their hidden hosts and then parasitize them, usually by using olfactory semiochemicals from hosts (lavare and adults), host frass and symbiotic microorganisms in host galleries; visual signals from color contrast of plant surface; contact stimuli from characters of host physical defense; substrate vibrations produced by host feeding and movement; infrared radiation from host activities and metabolizability. Some parasitic wasps may integrate the information of several stimuli from different sources to enhance the reliability and accuracy of host locations. In addition, the potentials for utilizing the host location signals of parasitoids in biological control are discussed.     

Host-feeding and acceptance by a parasitic wasp (Hymenoptera: Ichneumonidae) as influenced by egg load and experience in a patch

Optimal host selection models based on dynamic programming predict that the physiological state of a foraging insect, i.e. egg load, energy reserves etc., influences behavioral decisions. To test this prediction, the effect of physiological state on host acceptance of the ectoparasitic wasp Agrothereutes lanceolatus was investigated. Female wasps in plastic cups (regarded as patches) were presented with hosts, and their responses to the hosts were continuously observed. After observations, the wasps were dissected and the number of mature and immature eggs they carried were counted. The results showed that behavioral decisions by the female wasps were influenced by mature egg load, but not by wasp size or immature egg load. Hence the wasps with higher egg loads were more likely to oviposit. The number of hosts previously encountered in a patch (i.e. wasp experience) also had an independent effect on females' host acceptance, indicating that female informational state was updated during foraging in that patch. Female wasps host-fed only when mature egg load approached zero. Concurrent host-feeding was not observed. Parasitoid survival was almost zero when parasitoid eggs were transferred onto hosts that were fed upon, indicating that concurrent host-feeding could cause a high degree of offspring mortality. These three results supported the assumption and prediction of optimal host-feeding models. Parasitoid host selection and host-feeding are discussed in the context of recent models.     

Host-size-dependent sex ratios among parasitoid wasps: does host growth matter?

Summary Waage's (1982) hypothesis that host-size-dependent sex ratios will occur in parasitoids of nongrowing hosts and not in parasitoids of growing hosts is examined using published data on parasitoid wasps. Waage's hypothesis is supported as a general, but not absolute, rule: among solitary parasitoid wasps, a significantly greater proportion of parasitoids of nongrowing than of growing hosts show some evidence of host-size-dependent sex ratios (85% versus 42%, G=6.54, P< 0.05). The premise of Waage's hypothesis-that for parasitoids which develop in a growing stage, host size at oviposition is not a good predictor of the amount of resources available to the developing parasitoid-is also examined. It is suggested that across host species Waage's premise will hold for some, but not all, parasitoids of growing hosts. Likely exceptions to Waage's premise, and thus his prediction, are discussed. Parasitoids of growing hosts which are expected to have evolved hostsize-dependent sex ratios include parasitoids which utilize a narrow size range of host species, parasitoids which can distinguish among host species by some criterion other than size, and parasitoids which utilize host species whose susceptible instars do not overlap in size.     

Shifting preference between oviposition vs. host-feeding under changing host densities in two aphelinid parasitoids

Destructive host-feeding is common in hymenopteran parasitoids. Such feeding may be restricted to host stages not preferred for oviposition. However, whether this is a fixed strategy or can vary according to resource levels or parasitoid needs is less clear. We tested the trade-off between host feeding and oviposition on two whitefly parasitoids under varying host densities. Females of two aphelinid parasitoids, Eretmocerus hayati and Encarsia sophia were exposed to nine different densities of their whitefly host, Bemisia tabaci, in single-instar tests to identify their functional response. Mixed-instar host choice tests were also conducted by exposing whiteflies at four densities to the parasitoids. We hypothesized that the parasitoid females can detect different host densities, and decide on oviposition vs. host-feeding accordingly. The results showed that both Er. hayati and En. sophia females tended to increase both oviposition and host-feeding with increased host density within a certain range. Oviposition reached a plateau at lower host density than host-feeding in Er. hayati, while En. sophia reached its oviposition plateau at higher densities. At low densities, Er. hayati parasitized most on first and second (the optimal ones), and fed most on third nymphal instars (the suboptimal one) of the whitefly host as theory predicts, while at high densities, both parasitism and host-feeding occurred on first and second instars which are preferred for oviposition. En. sophia parasitized most on third and fourth (the optimal ones), while fed on first instars (the suboptimal one) at low densities, and utilized third and fourth instars for both at high densities. In conclusion, oviposition vs. host-feeding strategy of parasitoid females was found to vary at different host densities. The balance between reserving optimal hosts for oviposition or using them for host-feeding depended on parasitoid life history and the availability of host resources.     

Adult diet affects the life history and host-killing behavior of a host-feeding parasitoid

Neochrysocharis formosa (Westwood), an important biocontrol agent of agromyzid leafminers worldwide, is a host-feeding, idiobiont parasitoid. Female wasps have three types of host-killing behaviors: reproductive (parasitism), non-reproductive host feeding (host feeding), and host stinging without oviposition or feeding (host stinging). In this study, we compared the life history and host-killing behaviors of female parasitoids under four adult diets: starvation, hosts only, hosts plus honey (10% w/v honey solution), and honey only. Furthermore, we analyzed the host-feeding and oviposition preferences of adult females in the hosts-only and hosts-plus-honey treatments. Female parasitoids feeding on hosts had significantly increased longevity, higher fecundity, more host-stinging events, and caused a higher total host mortality than parasitoids in the starvation treatment. The honey supplement significantly increased longevity, fecundity, host-stinging events, and total host mortality, as well as average daily fecundity, but did not alter host-feeding events, host-stinging events, or daily total host mortality. However, the honey supplement did reduce the number of daily host-feeding events and induced a shift toward oviposition. Finally, we found that the non-reproductive host killing caused by host feeding and host stinging enhanced the control potential of N. formosa. These results should contribute to a better understanding of the biocontrol efficiency of destructive host feeders.     

Food-deprived host-feeding parasitoids kill more pest insects

Some parasitoids not only parasitize their hosts but also destructively feed on them which could result in host mortality. We hypothesized that host-feeding parasitoids that are food-deprived before being released for biological control would destructively feed on more hosts, and an optimal duration of food deprivation of parasitoids would enhance host feeding without negatively affecting parasitism. We investigated the effects of food-deprivation durations (0, 3, 6, 10, 24 h and 20% honey solution offered), before being released, on the host mortality and parasitism by a destructive host-feeding parasitoid of whitefly nymphs, Encarsia sophia (Girault & Dodd) (Hymenoptera: Aphelinidae). In another experiment, we compared host feeding, parasitization and longevity throughout the lifespan between food-deprived and non-food-deprived parasitoids. Using fourth-instar Bemisia tabaci B biotype as the host, the responses of E. sophia differed significantly with the durations of food deprivation before they were exposed to their hosts. E. sophia adults food-deprived for 6 h killed more hosts through feeding than those that were food-deprived for 0, 3, 10 and 24 h or those that were fed with honey solution. Moreover, parasitoids that were food-deprived for 6 h parasitized more hosts than those held for other durations of food deprivation. Similar results on host-feeding were found when third-instar whitefly nymphs were used. We confirmed that E. sophia food-deprived for 6 h not only killed more whiteflies through host feeding, but also lived significantly longer and parasitized more hosts than the non-food-deprived ones throughout their lifespan. Our results could be extended to improve the effectiveness of augmentative biological control of pests through improved host feeding and parasitizing capacity by starving the parasitoids for a certain period of time before they will be used.