寄生蜂取食寄主特性及其在害虫生物防治中的作用

许多寄生性天敌昆虫的雌虫不仅寄生寄主, 而且还能取食寄主。在卵育型（synovigenic）寄生蜂类群中, 取食寄主行为是较为普遍的现象。本文综合近20年相关研究进展, 从寄生蜂类群、取食类型、生态学意义及影响因子等方面对寄生蜂的取食寄主行为进行了归纳总结。寄生蜂通过取食不仅可以杀死寄主, 直接起到控制害虫种群数量的作用, 还能通过取食策略为卵的成熟和再生提供营养来源, 对延长雌虫的寿命也有一定的帮助。对取食寄主行为的了解可为筛选优势寄生性天敌种类、评估寄生蜂在害虫生物防治中的作用提供重要信息。     

蚜虫寄主专化型及其成因

综述了蚜虫寄主专化型产生的可能原因。蚜虫寄主专化型的形成在生态学水平上主要与蚜虫对寄主的选择识别能力、天敌作用、其他共生（共存）生物作用、抗药性等有关;并且具有一定的遗传基础,主要表现在蚜虫体内酶系的变化、染色体变异、有性繁殖中同型交配行为及种群的遗传分化上。但是对某种寄主专化型蚜虫而言,其具体的成因尚不明确。     

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.     

天敌昆虫利用信息化学物质寻找寄主或猎物的研究进展

应用天敌昆虫控制害虫是传统生物防治中的主要内容。信息化学物质是天敌昆虫远距离向寄主或猎物栖息地进行定向,以及近距离对寄主或猎物进行定位所依赖的重要信号。综述了能够引诱天敌昆虫的信息化学物质的产生、分类、作用、异同及背景,并介绍了天敌昆虫对信息化学物质的行为反应和特异性。同时,对这类物质在生物防治中的应用及前景作了介绍与探讨。对不同营养层的种间化学通讯作为研究协同进化时的重要参考也作了简要介绍。     

昆虫内共生菌在昆虫防御中的作用

共生菌在昆虫体内的生长、繁殖过程中起着十分重要的作用,如营养功能、解毒作用、调控生殖及与寄主适应性等等。近些年,一些学者提出了共生菌在寄主抵御天敌等逆境的胁迫时也具有重要的作用。其在寄主防御中的作用可分为3类:(1)昆虫体内共生菌能产生具有抗真菌作用的代谢物,能够抵御病原真菌的入侵,提高寄主的适应性。(2)昆虫抵御寄生蜂的作用与其体内的共生细菌密切相关。(3)共生菌也能产生有毒物质保护寄主免于被捕食。昆虫共生菌在寄主防御中的作用机制除了产生抗菌物质和聚酮类等有毒物质外,还可能通过与水平传播的寄生物(horizontal transmission parasites,HTPs)竞争寄主资源来直接保护寄主。对共生菌的作用的深入研究将有可能进一步明确其功能,并加以利用。     

昆虫翅型分化的表型可塑性机制

翅多型现象在昆虫中广泛存在,是昆虫在飞行扩散和繁殖能力之间权衡的一种策略,对种群的环境适应性进化具有重要的意义。目前在植食性昆虫中研究较多,有关寄生蜂的翅型分化鲜见报道。综述了昆虫翅型分化的表型可塑性机制。遗传因素和环境因素均对昆虫翅的发育产生影响,基因型对翅型的决定具有显著作用,外界环境条件,包括温度、光周期、食物质量、自身密度、外源激素等因素对昆虫翅的发育也产生重要的调节作用,从而产生翅的非遗传多型性现象。此外,天敌的寄生或捕食作用可能会诱导某些昆虫的翅型产生隔代表型变化。对昆虫产生翅多型现象的生态学意义及其在生物进化过程中的作用进行了讨论,并探讨了寄生性昆虫翅型分化机制在生物防治上的可能应用途径。功能基因组学和表观遗传学的进一步发展可望为彻底揭示昆虫翅型分化机制提供新的机遇和技术手段。     

二化螟寄主种群及种群间生殖隔离形成机制研究

水稻和茭白是二化螟的两种主要寄主植物。经过长期的适应进化,二化螟已分化出水稻种群和茭白种群。国内外的大量研究表明,二化螟水稻种群和茭白种群在形态特征、交配行为、寄主选择性、寄主适应性、田间种群动态、越冬生物学、生理生化及遗传多样性等方面均存在一定差异。二化螟水稻种群和茭白种群的发生期差异、交配节律差异和交配后配子不亲合等可能造成了寄主种群的部分生殖隔离。目前认为,二化螟两寄主种群间仍存在基因交流,处于物种形成的早期阶段,但尚需从寄主种群鉴定、表型差异区分和种群间生殖隔离机制等方面开展深入研究来进一步明确,为理解昆虫同域物种形成机制提供参考。     

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。     

自复寄生蜂的生殖特性及其对生物防治的影响

自复寄生蜂营雌雄异律发育,雌蜂为初级寄生蜂,雄蜂为复寄生蜂。依据寄主类型产下的相应性别的后代,其性比受到寄主数量、初级寄主与二级寄主的相对丰度等因素影响。自复寄生蜂能够寄生和取食初级寄主,抑制有害昆虫的种群数量,同时能够寄生和取食同种和异种寄生蜂幼虫,产生致死干扰竞争作用。复寄生二级寄主时,易感时间窗口主要集中在老熟幼虫至预蛹阶段。与同种二级寄主相比,自复寄生蜂更倾向于寄生异种二级寄主或无选择倾向性,取食时,更倾向于异种二级寄主。自复寄生蜂特殊的生殖方式使其对生物防治的影响备受争议。在评价其生物防治价值时,应对其正面影响和负面影响做出全面权衡。本文对自复寄生蜂的生殖特性、对同种和其他种类寄生蜂产生的致死干扰竞争作用及其对生物防治的影响等问题的国内外研究进展进行了综述,以期为利用这种天敌资源开展有害生物综合治理提供依据。     

我国烟粉虱主要捕食和寄生性天敌控制能力研究进展

烟粉虱Bemisia tabaci Gennadius是一种广泛分布于世界各地的多食性害虫,对我国多个省市区的农作物和观赏植物造成了严重的危害。化学杀虫剂的大量使用在杀伤天敌的同时使烟粉虱产生了不同程度的抗性,防治越来越困难;相较于化学防治,在不同地区筛选有效的天敌对烟粉虱进行生物防治是对烟粉虱进行有效控制的重要方法。我国报道的捕食性天敌共计26科109种,寄生性天敌共计2科59种。烟粉虱防治过程中由于单一的防治效果不理想,联合多个天敌资源的利用是增加对烟粉虱的生物防治效果的重要手段。本文通过对我国烟粉虱捕食性和寄生性天敌种类系统全面的整理,同时对烟粉虱优势种天敌中的单种或者多种组合利用后对烟粉虱的的控害潜能进行综述,以期为烟粉虱天敌昆虫种类的筛选和实现高效的生物防治提供科学依据。     

Income resources and reproductive opportunities change life history traits and the egg/time limitation trade‐off in a synovigenic parasitoid

1. Synovigenic parasitoids emerging with no or only a few mature oocytes could not rely on only capital resources, but also need to acquire income resources. Income resources in nature can either contribute to egg maturation as a food resource and/or create unpredictability in realised reproductive opportunities for synovigenic parasitoids. Therefore, we hypothesised such resources could affect life history traits and the risks of egg/time limitation in synovigenic parasitoids. 2. Using the Ovigeny Index, we investigated the effects of various host availability levels (unavailable, limited, and unlimited availability) and non‐host foods (water and honey) on life history traits and on the occurrence of egg/time limitation in Eretmocerus hayati, a predominant parasitoid on Bemisia tabaci. 3. The Ovigeny Index of Er. hayati was 0.28, which suggested it was a typical synovigenic species. Both host availability levels and non‐food type had major effects on life history traits of this parasitoid, but the availability of hosts for both feeding and reproduction was the key factor. Meanwhile, egg/time limitation was encountered by all wasps and its intensity varied with host availability levels. 4. Our results confirmed that the income resource and reproductive opportunity played a central role in shaping the life history and risks of egg/time limitation of a synovigenic parasitoid.     

Differential attraction of parasitoids in relation to specificity of kairomones from herbivores and their by‐products

Infochemicals are used by foraging parasitoids in the host selection process from habitat preference until host recognition. Kairomones from the herbivore host plays a vital role in the attraction of parasitoids, particularly in the micro‐habitat. Parasitoids are specifically attracted to their respective herbivore species even when different herbivores are present on the same plant. Chemicals emitted from different stages of host (eggs, larvae, pupae, adult), host by‐products (e.g., frass, exuviae, mandibular gland secretions, defense secretions etc.), or intra‐specific infochemicals (pheromones) can be main signals for the parasitoids. Parasitoids can differentiate between host and non‐host, between different hosts and host stages by perceiving specific volatile and contact kairomones from the host itself, host along with its by‐product, by‐products alone or intra‐specific infochemicals; of which frass (by‐product) and intra‐specific infochemicals are the most reported ones. Adult and larval parasitoids have been reported to be attracted to kairomones of their target stage or byproduct of their host. Pupal parasitoids have been found to utilize kairomones from the preceding host stage while egg parasitoids are known to exploit a variety of host infochemicals, for example, either from eggs themselves or other non‐target host stages, especially adults and adult‐related by‐products. The kairomonal chemicals identified so far include various groups, but mainly hydrocarbons. A high degree of host specificity and host acceptance is important for the parasitoids as any mistake may result in the loss of fitness.     

Odour-mediated long-range avoidance of interspecific competition by a solitary endoparasitoid: a time-saving foraging strategy

1. In studies on optimal foraging strategies, long-range decisions in the pursuit of resource are rarely considered. This is also the case for sympatric parasitoids, which may be confronted with the decision to accept or reject host larvae that are already parasitized by a competing species. They can be expected to reject already parasitized hosts if it is likely that they will lose the resulting intrinsic competition. However, examples of such interspecific host discrimination are rare. 2. We propose that parasitoids that are not egg-limited should reject inferior hosts only if it saves them time, and that this will be achieved mainly when the parasitoids are able to detect competitors from a distance. We tested this hypothesis using the sympatric parasitoids Cotesia marginiventris (Cresson) and Campoletis sonorensis (Cameron). 3. C. sonorensis was found to be the superior intrinsic competitor but, upon contact with a host larva, both wasps readily accepted hosts that had already been parasitized by the other species. However, in an olfactometer experiment, C. marginiventris females were found to strongly avoid the odour of their superior competitor. 4. These results are in accordance with a time optimization scenario, whereby the inferior competitor accepts competition if it costs only an egg, but avoids competition if it may save time that can be allocated to the search for more profitable hosts. 5. Models on host discrimination strategies in parasitoids had not yet considered discrimination from a distance. Long-range foraging decisions can also be expected for other organisms that have to choose between resources of varying suitability and profitability.     

补充非寄主食物对芙新姬小蜂和潜蝇姬小蜂控害行为权衡的影响

芙新姬小蜂和潜蝇姬小蜂均为强卵育型(synovigenic)寄生蜂,成虫具有产卵寄生致死、寄主取食致死和寄主叮蛰直接致死3种寄主致死行为。为了明确成虫补充非寄主食物营养对2种姬小蜂控害潜力/控害行为的影响,本文比较研究了分别提供寄主食物和寄主食物+非寄主食物(10%葡萄糖)条件下的2种姬小蜂在初羽化前期(前6 d)内对美洲斑潜蝇幼虫的致死行为的影响。结果显示:在仅提供寄主食物时,芙新姬小蜂的寄生、取食和总体致死量均显著的高于潜蝇姬小蜂,但是直接致死能力却弱于后者;补充非寄主食物和仅提供寄主食物处理相比,两种寄生蜂的寄生量,取食量和总致死量均呈下降趋势。该研究显示:在初羽化前期,取食寄主食物的雌蜂比取食寄主食物+葡萄糖的雌蜂具有更强的致死能力。     

Cascading effects of host size and host plant species on parasitoid resource allocation

1. The bottom‐up factors that determine parasitoid host use are an important area of research in insect ecology. Host size is likely to be a primary cue for foraging parasitoids due to its potential influence on offspring development time, the risk of multiparasitism, and host immunocompetence. Host size is mediated in part by host‐plant traits that influence herbivore growth and potentially affect a herbivore's quality as a host for parasitoids. 2. Here, we tested how caterpillar host size and host plant species influence adult fly parasitoid size and whether host size influences wasp parasitoid sex allocation. We measured the hind tibia lengths and determined the sex of wasp and fly parasitoids reared from 11 common host species of polyphagous caterpillars (Limacodidae) that were in turn reared on foliage of seven different host plant species. 3. We also tested how host caterpillar species, host caterpillar size, and host and parasitoid phenology affect how the parasitoid community partitions host resources. We found evidence that parasitoids primarily partition their shared hosts based on size, but not by host species or phenology. One index of specialisation (d′) supports our observation that these parasitoids are quite generalised within the Limacodidae. In general, wasps were reared from caterpillars collected in early instars, while flies were reared from caterpillars collected in late instars. Furthermore, for at least one species of solitary wasp, host size influenced sex allocation of offspring by ovipositing females. 4. Host‐plant quality indirectly affected the size attained by a tachinid fly parasitoid through its direct effects on the size and performance of the caterpillar host. The host plants that resulted in the highest caterpillar host performance in the absence of enemies also yielded the largest parasitoid flies, which suggests that host plant quality can cascade up to influence the third trophic level.     

半闭弯尾姬蜂寄主搜索中的学习行为

研究了半闭弯尾姬蜂寄主搜索过程中的学习行为。结果表明,成虫期之前的饲养寄主所取食的寄主植物对成蜂行为没有影响,而雌蜂早期的短暂经历可对其随后的行为反应产生显著影响,从而对已经历的植物气味表现出显著的嗜好,但这种通过学习所表现出的嗜好又可因新的经历而改变。雌成蜂不仅能对其所经历的虫伤寄主植物释放的信息化合物进行学习,而且对其所经历的寄主幼虫的信息化合物也能进行学习。     

Factors affecting the evolution of development strategies in parasitoid wasps: the importance of functional constraints and incorporating complexity

Parasitoid wasps have long been considered as model organisms for examining optimal resource allocation to different fitness functions, such as body size and development time. Unlike insect predators, which may need to consume many prey items to attain maturity, parasitoids generally rely on a limited amount of resources that are obtained from a single source (the host). This review discusses a range of ecophysiological constraints that affect host quality and concomitantly the evolution of development strategies in parasitoids. Two macroevolutionary differences in host usage strategies (idiobiosis, koinobiosis) are initially described. Over many years, particular attention has been paid in examining a range of quantitative host attributes such as size, age, or stage, as these affect idiobiont and koinobiont parasitoid development. Parasitoids and their hosts, however, constitute only a small part of an ecological community. Consequently, host quality may be affected by a broad range of factors that may operate over variable spatial and temporal scales. Intimate factors include aggressive competition with other parasitoids and pathogens for access to host resources, whereas less intimate factors include the effects of toxic plant compounds (allelochemicals) on parasitoid performance as mediated through primary and/or secondary hosts. It is suggested that future experiments should increase the levels of trophic complexity as these influence the evolution of life history and development strategies in parasitoids. This includes integration of a suite of direct and indirect mechanisms, including biological processes occurring in different ecological realms, such as above‐ground and below‐ground interactions.     

昆虫寄主标记信息素

综述了昆虫寄主标记信息素的研究进展。昆虫寄主标记信息素是指由昆虫产生的用来标记寄主上有同种个体存在的化学物质。昆虫寄主标记信息素的主要生态学功能是调节昆虫的产卵行为,通过阻止自身或同种其它个体对已标记寄主的产卵选择,或减少产卵量来减少后代之间对寄主资源的竞争。寄主标记信息素也会给释放着带来不利的影响,如信息盗用和盗寄生现象等。昆虫寄主标记信息素也调节昆虫近缘种之间对共同寄主资源的竞争。近缘种昆虫对相互寄主标记信息素识别能力的差异反映了不同昆虫对同一寄主资源竞争能力的强弱。寄主标记信息素产生和贮存的部位一般与外分泌腺、消化系统或生殖系统相联系,杜氏腺、毒腺、前胸腺、腹腺、下唇腺、后产卵管、卵巢、中肠和后肠等是产生或贮存寄主标记信息素的常见部位。产生的寄主标记信息素一般在成虫产卵时由产卵器、口器或排泄口释放到寄主体内或体表。卵寄生蜂的寄主标记信息素一般标记在寄主的体表,雌成蜂用触角检测;其它寄生蜂的寄主标记信息素常产在寄主体内,用产卵器检测;植食性昆虫的寄主标记信息素只产在寄主表面,用触角或产卵器检测。昆虫的产卵器、口器、触角或跗节上着生有感受寄主标记信息素的化感器,可以检测到标记在寄主体内或体表的寄主标记信息素。昆虫寄主标记信息素的完全定性涉及活性化合物的分离、鉴定、合成以及行为测定等,已有几种昆虫的寄主标记信息素成分得到了分离鉴定。     

Comparative phylogeography across two trophic levels: the oak gall wasp Andricus kollari and its chalcid parasitoid Megastigmus stigmatizans

Insect parasitoids are important components of many terrestrial ecosystems. However, relatively little is known about the mechanisms responsible for structuring their populations. Here we investigate the ability of Megastigmus stigmatizans, an oak gall wasp parasitoid, to track its host Andricus kollari over two different timescales, and examine its current population structure across a divide in host population structure. The divide represents a transition in gall wasp host-plant species and offers the opportunity to examine whether the split, which divides gall wasp populations, manifests itself in the next trophic level. Analysis of mitochondrial haplotype data for parasitoid and host reveals: (i) A similar phylogeographic population structure for both, with Iberian populations more derived with respect to more eastern populations. (ii) It is likely that the host colonized the Iberian refuge earlier than the parasitoid, probably by at least one glacial cycle. (iii) Recent range expansion of central European host populations northwards has resulted in pursuit by parasitoids from the same geographic origin. (iv) In addition, Iberian parasitoid populations have crossed a major divide in host population structure to invade northern Europe. Such human-facilitated escape from natural refugial distributions may have important implications for the composition and structure of northern European gall wasp communities.     

Does mother really know best? Oviposition preference reduces reproductive performance in the generalist parasitoid Aphidius ervi

The reproductive success of female parasitoids is dependent on their ability to accurately assess the suitability of a host for larval development. For generalist parasitoids, which utilize a broad range of species and instars as hosts, a set of assessment criteria determines whether a host is accepted or rejected. The suitability of a host, however, can only be imperfectly assessed by the female parasitoid, which can result in the selection of lesser quality hosts for oviposition. In this study we explored the disparity between host quality and host preference using the generalist koinobiotic parasitoid Aphidius ervi Haliday (Hymenoptera: Aphidiidae) and the host Aulacorthum solani (Harris) (Homoptera: Aphididae), the foxglove aphid. The second instar hosts produced the highest level of reproductive success, while third and fourth instars resulted in a substantially reduced reproductive performance. When given a choice of host instars, parasitoids preferred the older hosts for oviposition disregarding their reduced suitability for larval development. Results are discussed in context of mechanisms involved in A. ervi host selection and biases in the criteria used to assess hosts that may arise when parasitoids transfer host species between generations.