虫瘿与致瘿昆虫

虫瘿是由昆虫等致瘿生物诱导寄主植物而产生的一种特异组织。虫瘿对致瘿生物具有提供营养和保护等作用,虫瘿不仅有一定的经济利用和科学研究价值,而且多数致瘿昆虫是农林害虫。本文对致瘿昆虫在植物上的致瘿部位、致瘿的主要昆虫类群、虫瘿形态结构、虫瘿化学组成、虫瘿对寄主植物的影响以及虫瘿的利用等进行综述。     

虫瘿多样性及其与寄主植物和环境间关系

虫瘿是自然界极常见的生物现象,凝聚着昆虫与植物间显著、复杂而密切的协同关系。本文主要阐述了致瘿昆虫的主要类群及其在植物上的致瘿部位、虫瘿外部形态、虫瘿发育过程、虫瘿内部结构、虫瘿寄主植物多样性以及虫瘿空间分布规律等,探讨了致瘿昆虫和寄主植物间相互关系,以及影响虫瘿空间分布的环境因素等。最后对目前虫瘿生物学存在的问题及以后的研究方向进行了讨论,以期为有害虫瘿的控制和有益虫瘿的开发与利用,以及致瘿昆虫与寄主植物间协同演化关系、致瘿昆虫的致瘿机理等研究奠定一定的理论基础。     

虫瘿挥发物对枸杞红瘿蚊拟长尾小蜂寄主选择的影响

[目的]为明确枸杞红瘿蚊拟长尾小蜂Pseudotorymus jaapiellae Yang et Chen对枸杞红瘿蚊Gephyraulus lycantha Jiao&Kolesik诱导产生虫瘿挥发物的选择差异性,解析影响其寄主选择的信息化学物质.[方法]采用Y型嗅觉仪,测定枸杞红瘿蚊拟长尾小蜂的孕卵雌蜂对受枸杞红瘿蚊为害的幼嫩虫瘿和成熟虫瘿挥发物的行为选择偏好;通过顶空固相微萃取和气相色谱-质谱联用技术,分析影响该寄生蜂产卵选择的不同发育阶段虫瘿挥发物成分变化与差异.[结果]与枸杞健康花蕾相比,枸杞红瘿蚊拟长尾小蜂明显趋向于选择受枸杞红瘿蚊为害的虫瘿,尤其是幼嫩虫瘿对孕卵雌蜂具有更显著的吸引作用(P＜0.01),其中有95％的寄生蜂选择幼嫩虫瘿,仅有5％的寄生蜂选择健康花蕾;不同发育阶段虫瘿与健康花蕾的挥发物种类和释放量存在明显差异,幼嫩虫瘿的挥发物种类最多,达61种,成熟虫瘿和健康花蕾分别为52种和54种;酮类和萜类化合物是枸杞花蕾受害后新出现的物质,其中萜类化合物仅存在于幼嫩虫瘿中,酮类化合物随着虫瘿的成熟,其含量逐渐升高.[结论]枸杞红瘿蚊拟长尾小蜂能够通过对枸杞不同发育阶段受害虫瘿的特异性挥发物进行寄主识别,从而定位寄主,实现对害虫枸杞红瘿蚊的种群调控.     

5种生物杀虫剂对4种天敌昆虫的安全性评价

【目的】生物杀虫剂及天敌昆虫的应用是绿色防控的发展方向,但有关生物杀虫剂对天敌昆虫的安全性尚不明确。研究生物杀虫剂对天敌昆虫的影响可以为更好地协调使用生物杀虫剂和天敌昆虫提供理论依据。【方法】在实验室条件下采用药膜法和滤纸膜片法测定了5种生物杀虫剂(苦参碱、桉油精、鱼藤酮、除虫菊素、橄榄鲨)在田间推荐使用浓度下,对4种天敌昆虫(丽蚜小蜂、东亚小花蝽、食蚜瘿蚊、巴氏新小绥螨)的致死率。【结果】除虫菊素对丽蚜小蜂、东亚小花蝽、巴氏新小绥螨的影响较大,致死率均达到100%;鱼藤酮对丽蚜小蜂、食蚜瘿蚊、巴氏新小绥螨的影响均较大,致死率在98.65%以上;橄榄鲨对丽蚜小蜂、东亚小花蝽、食蚜瘿蚊的影响较小,致死率均小于30%;苦参碱对丽蚜小蜂的影响最大,致死率达100%,对食蚜瘿蚊和巴氏新小绥螨的影响较小,致死率分别为15.56%和15.91%;桉油精对巴氏新小绥螨的影响最大,致死率达100%,对东亚小花蝽和食蚜瘿蚊的影响较小,致死率分别为15.91%和6.67%。【结论】这5种生物杀虫剂中,橄榄鲨对丽蚜小蜂、东亚小花蝽、食蚜瘿蚊安全,桉油精对东亚小花蝽、食蚜瘿蚊安全,苦参碱对食蚜瘿蚊、巴氏新小绥螨安全,鱼藤酮对东亚小花蝽安全,除虫菊素对食蚜瘿蚊安全。     

虫瘿与其生物群落及寄主植物间的关系

虫瘿是自然界极其常见的生物现象,是植物与昆虫互作的奇特产物。本文对虫瘿生物群落多样性、虫瘿与其生物群落的关系以及虫瘿与寄主植物的关系进行概述,探讨了致瘿昆虫在虫瘿形成中的作用、植物化学对致瘿昆虫产卵交配行为的影响以及植物激素在虫瘿形成中的作用,最后对虫瘿今后的研究方向进行了讨论,为虫瘿的致瘿生物学及其瘿内生物相互关系的进一步研究奠定基础。     

一种新的危害芒果树叶的瘿蚊害虫

瘿蚊科中有些种类是危害芒果的重要害虫 ,全世界已报道危害芒果的瘿蚊约有 1 0属 ,2 0多种。中国有 6种 ,其中危害芒果树叶片的占 3种。近几年在厦门地区发现芒果树叶片被寄生大量虫瘿 ,导致叶片卷缩、枯萎、脱落。笔者在厦门采集到大量受害芒果树叶片上的虫瘿 ,解剖到了幼虫、蛹 ,并饲养出了成虫。经研究 ,确认该瘿蚊为普瘿蚊属 (中国新记录属 )的一个新种 ,定名为壮铗普瘿蚊ProcontariniarobustaLi,BuandZhang ,2 0 0 3[1] ,于 2 0 0 3年发表在《动物分类学报》第 2 8卷第 1期。现将该虫的鉴别特征及其在厦门地区的发生、分布和为害情况…     

食蚜瘿蚊全虫态形态记述

【目的】基于实验室饲养的食蚜瘿蚊Aphidoletes aphidimyza (Rondani)进行个体发育的形态特征描述。为食蚜瘿蚊的鉴定提供基本信息。【方法】在野外采集幼虫,温度(25±1)℃、相对湿度65%±5%、光照16 L∶8 D的室内条件下饲养获得食蚜瘿蚊卵、幼虫、蛹和成虫的全部虫态。通过数码采集和电子显微镜详细描述了各虫态的发育特征。【结果】食蚜瘿蚊卵的发育历期为2 d,幼虫期有3个龄期,发育历期5-8 d,蛹期7-10 d以及成虫期3-4 d。【结论】食蚜瘿蚊个体发育特征对食蚜瘿蚊的鉴定和分类有指导意义。     

中国垫瘿蚊属研究(双翅目：瘿蚊科)

记述了双翅目瘿蚊科一中国新记录属垫瘿蚊属 Conarete Pritchard 的4 种,其中包括2新种： 短须垫瘿蚊 C. brevipalpa Li et Bu, sp. Nov.（模式产地： 海南尖峰岭）和葫茎垫瘿蚊 C. sicyoidea Li et Bu, sp. Nov.（模式产地：海南尖峰岭）; 另外2种为在中国首次记录的种： 米垫瘿蚊 C. mihijamensis Grover （分布：海南尖峰岭）和印垫瘿蚊 C. indorensis Grover（分布：陕西周至板房子）。文中给出了垫瘿蚊属与近缘的短角瘿蚊属Anarete Haliday的区别特征和中国种类分种检索表。模式标本均保存于南开大学生物学系昆虫标本室。     

温度对食蚜瘿蚊生长发育的影响

在19,22,25,28,31℃和RH为80%的组合下,测定食蚜瘿蚊Aphidoletes aphidimyza Rondani的发育历期,分析发育速率与温度的关系,并且测定各温度下食蚜瘿蚊的化蛹率和羽化率等生物学参数。实验结果表明:温度对食蚜瘿蚊的生长发育有较大影响。在19~28℃范围内,食蚜瘿蚊各虫态的发育历期随温度的升高而缩短,而在28~31℃范围内,食蚜瘿蚊各虫态的发育历期随着温度的升高而略为延长。采用线性日度模型和Logistic模型对卵期、幼虫期、蛹期和全世代的发育速率进行模拟分析,2种模型均能较好地反映各虫态的发育速率。不同温度下食蚜瘿蚊的5天化蛹率和总羽化率差别较大,但以25℃下为最高,分别为88.00%和94.70%;而22℃下的化蛹率和羽化率与25℃下较为接近,分别为84.00%和90.66%,经分析差异不显著。食蚜瘿蚊最适生长发育温度为22~25℃。     

不同冷藏条件对食蚜瘿蚊过冷却点和冰点的影响

【目的】食蚜瘿蚊Aphidoletes aphidimyza(Rondani)是一种捕食性天敌,其幼虫对蚜虫具有较好的控制潜能。研究不同冷藏条件对食蚜瘿蚊耐寒能力的影响,为食蚜瘿蚊的低温冷藏技术提供了理论依据。【方法】利用热电偶原理,测定食蚜瘿蚊在不同冷藏条件下的过冷却点和结冰点。【结果】25℃条件下,食蚜瘿蚊各虫态的过冷却点及结冰点存在显著差异（P<0.05）,其中2龄幼虫过冷却点（﹣26.18℃）和结冰点（﹣24.98℃）最低,雄成虫过冷却点（﹣22.95℃）和结冰点（﹣21.86℃）最高。在不同变温条件下冷藏食蚜瘿蚊蛹,其过冷却点在冷藏10、20和30 d时均上升且高于对照,冷藏40 d时低于对照;同一冷藏时长下不同冷藏方式间过冷却点的差异均不显著（P>0.05）。食蚜瘿蚊蛹在5℃条件下冷藏50d,每天定时7℃中断4h的结冰点（﹣23.09℃）最低,冷藏30d的结冰点（﹣21.15℃）最高;食蚜瘿蚊蛹在5℃条件下冷藏40 d,每天定时9℃中断4 h的结冰点（﹣22.66℃）最低,冷藏20 d时的结冰点（﹣20.95℃）最高;同一冷藏时长下3种冷藏方式间结冰点差异均不显著（P>0.05）。【结论】食蚜瘿蚊2龄幼虫的耐寒能力最强,雌成虫的耐寒能力要高于雄成虫;变温贮存和恒定低温均适合食蚜瘿蚊的低温贮存。     

Fungal endophytes which invade insect galls: insect pathogens,benign saprophytes,or fungal inquilines?

Fungi are frequently found within insect galls. However, the origin of these fungi, whether they are acting as pathogens, saprophytes invading already dead galls, or fungal inquilines which invade the gall but kill the gall maker by indirect means, is rarely investigated. A pathogenic role for these fungi is usually inferred but never tested. I chose the following leaf-galling-insect/host-plant pairs (1) a cynipid which forms two-chambered galls on the veins of Oregon white oak, (2) a cynipid which forms single-chambered galls on California coast live oak, and (3) an aphid which forms galls on narrowleaf cottonwood leaves. All pairs were reported to have fungi associated with dead insects inside the gall. These fungi were cultured and identified. For the two cynipids, all fungi found inside the galls were also present in the leaves as fungal endophytes. The cottonwood leaves examined did not harbor fungal endophytes. For the cynipid on Oregon white oak, the fungal endophyte grows from the leaf into the gall and infects all gall tissue but does not directly kill the gall maker. The insect dies as a result of the gall tissue dying from fungal infection. Therefore, the fungus acts as an inquiline. Approximately 12.5% of these galls die as a result of invasion by the fungal endophyte.     

The number of cecidomyiid insect galls affects the photosynthesis of Machilus thunbergii host leaves

Previous studies of the impacts of galls on host leaf photosynthesis do not suggest any general trends, with a reported range of effects from negative to positive. In this study, photosynthetic characteristics such as chlorophyll fluorescence (Fv/Fm), photosynthetic capacity, and stomata conductance were determined in two types of fruit-like galls (red ovoid and green obovate galls) induced by Daphnephila taiwanensis and Daphnephila sueyenae, respectively, in order to investigate whether the number of galls affects the photosynthesis of galled leaves of Machilus thunbergii. In 2008, chlorophyll fluorescence and photosynthetic capacity were negatively correlated with gall numbers, non-significantly and significantly, respectively, whereas stomata conductance was positively but non-significantly correlated with gall numbers. In 2009, photosynthesis capacity and stomata conductance were negatively, but non-significantly, correlated with gall numbers. Results imply that photosynthesis in M. thunbergii leaves is slightly affected by the number of cecidomyiid insect galls, and that the higher the gall number, the greater the negative effect that galls have on host leaf photosynthesis and subsequent infection.     

Plant organ abscission and the green island effect caused by gallmidges (Cecidomyiidae) on tropical trees

Plants exhibit a wide array of inert and induced responses in defense against herbivore attack. Among these the abscission of organs has been argued to be a highly effective mechanism, depending, however, on the herbivore’s feeding mode. While consisting of plant tissues, insect induced galls are seen as the extended phenotype of the gall inducer which might circumvent many or most of the plant defenses. There is very little information whether and how far beyond the gall tissue gall inducers might affect plant tissues. A localized impact is likely to leave the abscission of galled organs as a viable defense although at a cost. Here, we report on an instance where the host plant, Neea madeirana (Nyctaginaceae) abscises leaves galled by two species of Bruggmannia (Diptera: Cecidomyiidae), more frequently than ungalled leaves in a rain forest in Amazonia, Brazil. Once on the forest floor the leaves decay quickly, while both gall types show signs of localized maintenance of healthy tissues for a while (the green island effect). However, on the forest floor galls are exposed to a new set of potential natural enemies. Both gall types show a minimum of a five-fold increase in mortality due to pathogens (fungi and bacteria) compared to galls that were retained on the host tree. We discuss the adaptive nature of plant organ abscission as a plant defense against gallers and as a gall inducer adaptive trait. Handling editor: Graham Stone.     

Histocytological aspects of four types of ambrosia galls on Machilus zuihoensis Hayata (Lauraceae)

Four types of prosoplasmatic galls induced by Daphnephila midges are found on leaves of Machilus zuihoensis, a species endemic to Taiwan: urn- and small urn-shaped, obovate, and hairy oblong galls. In addition to containing nutritive tissues, these galls are lined with fungal hyphae. The objective of this study was to describe and compare the structural organization of the various gall morphologies and to examine the ultrastructure of the nutritive and fungal cells lining the gall chambers. The morphology and ultrastructure of mature-stage galls were examined by light, scanning electron, and transmission electron microscopy. Diverse epidermal cell shapes and wax textures were observed in the leaves and galls of M. zuihoensis. In small urn-shaped, obovate, and hairy oblong galls vascular bundles extend from the gall base to near the centre of the gall top. In contrast, vascular bundles in urn-shaped galls are distributed in the gall wall and extend to close to the outer gall top. Trichomes were present only abaxially on leaves and on hairy oblong gall surfaces. Starch granules, tannins, and mucilage were distributed differently among the four gall types. Further, fungal mycelia spread in the interior gall wall and partially passed through the intercellular spaces of nutritive cells and reached the sclerenchyma. Histological analyses revealed that the surface structure of galls differs from that of the leaf and that the epidermal organization differs among the four gall types. Different types of leaf galls on the same plant have different patterns of tissue stratification and contain different ergastic substances. The results of this study will contribute to the understanding of tritrophic relationships and the complex interactions among parasitic gall-inducing insects, mutualistic fungi, and host plants.     

Fungal diversity in galls of baldcypress trees

The baldcypress midge (Taxodiomyia cupressi and Taxodiomyia cupressiananassa) forms a gall that originates from leaf tissue. Female insects may inoculate galls with fungi during oviposition, or endophytes from the leaf tissue may grow into the gall interior. We investigated fungal diversity inside of baldcypress galls, comparing the gall communities to leaves and comparing fungal communities in galls that had successful emergence versus no emergence of midges or parasitoids. Galls of midges that successfully emerged were associated with diverse gall fungal communities, some of which were the same as the fungi found in surrounding leaves. Galls with no insect emergence were characterized by relatively low fungal diversity.     

Gall‐forming insect species richness along a non‐scleromorphic vegetation rainfall gradient in South Africa: The importance of plant community composition

Abstract Currently there is no single accepted hypothesis to explain gall‐forming insect species richness at a particular locality. Hygrothermal stress, soil nutrient availability, plant species richness, plant structural complexity, plant family or genus size, and host plant geographical range size have all been implicated in the determination of gall‐forming insect species richness. Previous studies of such richness at xeric sites have included predominantly scleromorphic vegetation, usually on nutrient‐poor soils. This study is the first to investigate gall‐forming insect species richness of xeric, non‐scleromorphic vegetation. Two habitat types were sampled at each of five localities across a rainfall gradient in the savanna biome of South Africa. The habitat types differed with respect to plant species composition and topography. Gall‐forming insect species richness did not increase with increasing hygrothermal stress or decreasing soil fertility. Rather, gall‐forming insect species richness was largely dependent on the presence of Terminalia sericea as well as other members of the Combretaceae and Mimosaceae. Plots where all these taxa were present had the highest gall‐forming insect species richness, up to 15 species, whereas plots with none of these taxa had a maximum of four galling‐insect species. Despite herb, shrub and tree strata not differing in gall‐forming insect species richness, insect galls were more common on woody than non‐woody plants. Also, stem galls were more frequent than apical or leaf galls. An alternative hypothesis to explain local gall‐forming insect species richness is suggested: galling insects may preferentially select those plant species with characteristics such as chemical toxicity, mechanical strength, degree of lignification or longevity that can be manipulated to benefit the galler. Thus plant community composition should be considered when attempting to explain gall‐forming insect species richness patterns.     

Mine and gall predation as top down regulation in the plant–insect systems from the Cretaceous of Negev, Israel

Fossil evidence of predation on leaf mines and galls opens the way to analyzing regulatory circuits in plant–insect interaction systems and assessing their evolutionary advancement. In the Cretaceous (mid-Turonian) flora of Negev, Israel, predation traces vary from the entire gall or mine excisions to minute punctures and slits over the mine tracks. Bite marks on the borders of predation holes representing different mouthpart morphologies may show how diverse the predators were. The efficiency of predation as a top down regulation force is attested on the basis of the gall and mine abundance, gall morphologies, mine configurations, co-occurrence of different mine types on leaves, and temporary mining. In turn, the regulation efficiency is considered as a criterion of evolutionary advancement of the plant–insect community as a whole. It is suggested that the coeval Cretaceous coastal and inland communities differed in the relative significance of top-down regulation.     

肚倍蚜对高单宁环境的抗氧化反应研究

肚倍蚜Kaburagia rhusicola Takagi是我国的一种重要的资源昆虫,其瘿内生活环境较特殊(虫瘿单宁含量为70%)。为了解肚倍蚜适应高单宁环境生化机制,本研究以性母为对照,测定了瘿内不同时期肚倍蚜体内总抗氧化能力(T-AOC)以及超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)和谷胱甘肽S-转移酶(GST)的活性,分析了肚倍蚜体内的抗氧化系统对高单宁环境胁迫所作出的相关反应。研究表明,与对照相比,瘿内时期肚倍蚜体内的总抗氧化能力、CAT和POD活性均显著升高;SOD和GST活性除在有翅蚜阶段与对照无差异,在其他阶段的活性均显著高于对照。本研究验证了高单宁环境对肚倍蚜造成氧化胁迫作用,认为肚倍蚜体内的抗氧化能力和抗氧化酶活性提高是其对高单宁环境的一种适应。     

Aphid galls accumulate high concentrations of amino acids: a support for the nutrition hypothesis for gall formation

The nutrition hypothesis for the adaptive significance of insect gall formation postulates that galls accumulate higher concentrations of nutritive compounds than uninfested plant tissue, resulting in a high performance of the gall former. This hypothesis has been supported by some taxa of gall insects, but not by taxa such as cynipid wasps. Aphid galls are expected to require higher levels of nitrogen than other insects’ galls with a single inhabitant, because aphid galls are required to sustain a number of aphids reproducing parthenogenetically over two generations. The present study tested this hypothesis by evaluating aphid performance and amino acid concentration in phloem sap, using the aphid Rhopalosiphum insertum (Walker) (Homoptera: Aphididae), which establishes colonies on leaves of Sorbus commixta Hedlund or in galls of the aphid Sorbaphis chaetosiphon Shaposhnikov (Homoptera: Aphididae). We prepared the gall and non‐gall treatments on trees of S. commixta, in which R. insertum fundatrices were reared and allowed to reproduce. In S. chaetosiphon galls, R. insertum colonies propagated more rapidly, and the second generation grew larger and more fecund than on ungalled leaves. The amount of amino acids exuding from cut galled leaves was fivefold that in ungalled leaves; however, there was no significant difference in the amino acid composition between galled and ungalled leaves. In the intact leaves, total amino acid concentration in the phloem sap declined rapidly from late April to late May; however, the galls retained this high amino acid concentration in developing leaves for 1 month. These results indicate that the improved performance in R. insertum is ascribed to the increased concentration of amino acids in galled leaves. We suggest that S. chaetosiphon galls function to promote the breakdown of leaf protein, leading to an increased performance of gall‐inhabiting aphids.     

Hormaphis hamamelidis and gall size: a test of the plant vigor hypothesis

We tested the Plant Vigor Hypothesis by determining the distribution of galls formed on leaves of witch hazel, Hamamelis virginiana , by the aphid Hormaphis hamamelidis , and by determining various factors that affect the fecundity of the gall-forming fundatrices. We also studied the role of the fundatrix in host plant manipulation. While the mean number of galls per leaf was low, galls had an aggregated distribution among leaves. Among trees, the average number of galls per leaf was not related to the mean leaf size, contrary to the preference prediction of the Plant Vigor Hypothesis. While fundatrices preferred the distal leaves of buds, which grew more than the proximal leaves, being on distal leaves conferred no increase in fecundity for fundatrices, contrary to predictions of the Plant Vigor Hypothesis. Gall size was the factor that explained the largest proportion of variation in fundatrix fecundity; fundatrix size explained somewhat less of the variation. Also, gall position on the leaf, number of aphid galls on the leaf, and on which leaf of the bud the gall was located all played small, statistically significant roles in explaining fundatrix fecundity, but their effects were variable between experiments. Removal of fundatrices shortly after galls had enclosed them limited the growth of galls, indicating the role of the fundatrices in gall growth. We compare and contrast this system versus other gall-forming insects, as well as discuss the adaptive significance of the aphid manipulation of the host plant. Much of the data contradict predictions of the Plant Vigor Hypothesis, and we discuss how gall size, as a measure of plant growth caused by insect manipulation, explains the observed patterns.