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。     

致瘿昆虫对寄主植物生理和代谢的影响

虫瘿是致瘿昆虫刺激植物后诱导形成的畸形结构,是研究植物与昆虫协同进化的理想材料,同时致瘿昆虫通常还是重要的农林害虫。因此,研究致瘿昆虫对寄主植物的影响,一方面可进一步揭示致瘿昆虫与植物的关系,有助于揭示成瘿植物生长的一般过程;另一方面,了解成瘿植物对致瘿昆虫的响应有助于筛选植物抗性指标、抗性基因、敏感基因等,为抗性育种提供理论基础。本文主要围绕致瘿昆虫对寄主植物光合作用、生理和代谢活动的影响等进行了综述。致瘿昆虫普遍引起寄主植物光合色素减少、光合速率降低,虫瘿内部组织中糖类、氨基酸类等初生代谢物质含量增加,虫瘿外部组织中酚类、黄酮类等非挥发性和萜类为主的挥发性次生代谢物质含量增加,寄主植物POD和SOD等保护酶活性增强,以及IAA, SA和JA等植物激素含量提高。现有研究资料显示,致瘿昆虫对寄主植物生理和代谢影响的研究仍处于初级阶段,其影响机制还需要进一步研究。     

中华蚊母树虫瘿类型及其致瘿昆虫生活史初探

初步观察了中华蚊母树(Distylium chinense(Fr.)Diels)不同类型虫瘿的发生过程及致瘿昆虫生活史。结果表明,中华蚊母树的虫瘿有3种类型,分别为叶/枝上大型虫瘿、叶上泡状虫瘿和叶柄/果上球型虫瘿。三者均为单室封闭型,次生开口。经鉴定致瘿昆虫均为半翅目蚜科昆虫,其中叶上泡状虫瘿致瘿昆虫为蚊母新胸蚜Neothoracaphis yanonis Matsumura。该虫3月初在虫瘿内营孤雌生殖,繁殖2代,5月底前飞离出瘿。     

向川安瘿蜂寄生白栎虫瘿和成瘿枝的代谢组比较

虫瘿是植物受致瘿昆虫产卵和取食等刺激而异常生长形成的不正常组织。营养假说认为虫瘿组织营养物质的含量高于寄主植物的成瘿部位,而次生代谢产物的含量低于寄主植物的成瘿部位。向川安瘿蜂(Andricus mukaigawae)属膜翅目(Hymenoptera)瘿蜂科(Cynipidae),在白栎(Quercus fabri)枝条上形成虫瘿。本文基于非靶向代谢组,使用高效液相色谱-质谱检测、鉴定和比较向川安瘿蜂幼虫期虫瘿与寄主植物成瘿枝的代谢物。研究结果表明,向川安瘿蜂虫瘿和成瘿枝的代谢物组成存在差异;虫瘿的脂肪含量均高于成瘿枝,虫瘿的4种单宁和4种黄酮物质的含量均低于成瘿枝,这支持营养假说;虫瘿的氨基酸含量均低于成瘿枝,部分酚类物质的含量高于成瘿枝,这不支持营养假说。     

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.     

植物的虫瘿与成瘿昆虫

虫瘿是植物组织遭受昆虫取食或产卵的刺激后,细胞加速分裂和异常分化而长成的畸形瘤状物或突起,它们是昆虫生活的"房子".介绍了虫瘿的形态多样性和形成过程,成瘿昆虫的多样性、生活史、寄生、食性和寄食现象,成瘿昆虫与寄主植物的关系以及人类对虫瘿的利用.     

植食性瘿蚊化学通讯中的信息化学物质

植食性瘿蚊是重要的农林害虫,其幼虫危害寄主植物,造成植物细胞分裂或异常分化而产生组织畸形,形成虫瘿。该虫具有成虫体小、寿命极短的特性,成虫羽化后可迅速求偶、交配,并随即寻找寄主植物产卵。在植食性瘿蚊短暂的生活史中要完成种群繁衍,配偶选择和寄主定位起着至关重要的作用,而植食性瘿蚊自身产生的信息素、寄主或非寄主产生的他感化学物质在调节其选择配偶和寄主植物过程中起着关键性的化学通讯作用。本文综述了国内外有关植食性瘿蚊化学通讯中信息化学物质的研究进展,包括植食性瘿蚊性信息素的释放、提取、组分鉴定、行为生测,以及寄主植物挥发物引诱活性物质的鉴定、筛选和对植食性瘿蚊定向和产卵行为的影响等方面,并对植食性瘿蚊信息化学物质的进一步研究和应用前景进行了探讨,以期为利用信息化学物质监测和防治植食性瘿蚊提供参考。     

宁夏枸杞对枸杞瘿螨为害的内源激素响应及外源水杨酸对枸杞瘿螨的影响

为探索宁夏枸杞对枸杞瘿螨致瘿过程诱导的激素响应及外源水杨酸对枸杞瘿螨的影响,利用LC-MS/MS技术对枸杞瘿螨为害后宁夏枸杞叶片中水杨酸(SA)、茉莉酸(JA)、生长素(Auxin)和脱落酸(ABA)4种内源激素进行测定,并通过浸叶法研究外源SA对枸杞瘿螨侵染过程、虫瘿生长和种群增长的影响。结果表明: 枸杞瘿螨的致瘿行为可引发枸杞叶片中多种内源激素产生变化,其中SA和JA的变化最显著,分别是对照组含量的4.0和13.0倍,而Auxin和ABA的变化不显著;外源SA对枸杞瘿螨虫瘿生长有显著抑制作用,且随时间的延长,抑制作用增强,同时虫瘿内瘿螨种群数量受到显著抑制。枸杞瘿螨能够同时诱导枸杞产生SA和JA抗性,外源SA对枸杞瘿螨虫瘿生长和瘿螨种群数量均具有显著抑制作用,利用外源SA防治枸杞瘿螨具有一定的应用前景。     

Evidence for long-distance, chemical gall induction by an insect

Abstract We report that a chemical stimulus from a herbivore, a galling insect, changes plant morphology and physiology to benefit the herbivore. Previous studies could not determine whether insect galls are induced by mechanical or chemical stimuli because feeding and oviposition both occurred at the site of gall formation. We report that the mouthparts of a spruce‐galling insect, Adelges cooleyi, were inserted in stem phloem cells far from induced galls, that tissues between mouthparts and galls appeared normal, and that the ability to initiate galls was inversely correlated with distance from buds (potential gall sites). Thus the effects of chemical stimuli were unambiguously separated from any mechanical influence of probing stylets or ovipositors. Our results strongly suggest that galls were induced by a chemical stimulus transported to buds via vascular tissue and that its efficacy was dose‐dependent.     

Gall-inducing Pachypsylla celtidis (Psyllidae) infiltrate hackberry trees with high concentrations of phytohormones

Abstract

In order to identify phytohormones involved in the initiation and maintenance of galls on the hackberry tree, Celtis occidentalis (Ulmaceae), in the presence of the insect Pachypsylla celtidis (Psyllidae); endogenous levels of cytokinins (CKs) and abscisic acid (ABA) were measured in the tissues of leaves, galls, and larval insects using liquid-chromatography-tandem-mass-spectrometry. The CKs isopentenyl adenine, isopentenyl adenosine, trans-zeatin and cis-zeatin, were extremely concentrated in insect larvae compared to surrounding tissue of leaves and galls. ABA concentrations in the insects were also relatively high at about 3268 pmol?g-1 fresh mass – approximately 25 times higher than in leaves, and 17 times higher than in galls. This represents a novel case of high concentrations of ABA being found in a larval insect. These findings indicate that manipulation of CK and ABA are involved in the initiation and maintenance of hackberry galls in the presence of larval P. celtidis.     

The parasitoid community of <Emphasis Type="Italic">Andricus quercuscalifornicus</Emphasis> and its association with gall size,phenology, and location

Plant galls are preyed upon by a diverse group of parasitoids and inquilines, which utilize the gall, often at the cost of the gall inducer. This community of insects has been poorly described for most cynipid-induced galls on oaks in North America, despite the diversity of these galls. This study describes the natural history of a common oak apple gall (Andricus quercuscalifornicus [Cynipidae]) and its parasitoid and inquiline community. We surveyed the abundance and phenology of members of the insect community emerging from 1234 oak apple galls collected in California’s Central Valley and found that composition of the insect community varied with galls of different size, phenology, and location. The gall maker, A. quercuscalifornicus, most often reached maturity in larger galls that developed later in the season. The parasitoid Torymus californicus [Torymidae] was associated with smaller galls, and galls that developed late in the summer. The most common parasitoid, Baryscapus gigas [Eulophidae], was more abundant in galls that developed late in the summer, though the percentage of galls attacked remained constant throughout the season. A lepidopteran inquiline of the gall (Cydia latiferreana [Tortricidae] and its hymenopteran parasitoid (Bassus nucicola [Braconidae]) were associated with galls that developed early in the summer. Parasitoids and inquilines, in general, had a longer emergence period and diapause within the gall than the gall-inducer. The association of different parasite species with galls of different size and phenology suggests that different parasite species utilize galls with slight differences in traits.     

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.     

Life-history traits in insect inclusions associated with bamboo galls

We examined the life-history traits of insect inclusions in bamboo galls induced by the gall maker, Aiolomorphus rhopaloides Walker (Hymenoptera: Eurytomidae) in a bamboo stand. Eight hymenopteran and one dipteran species were detected using soft X-ray photography of the galls and insect emergence from the galls. Aiolomorphus rhopaloides was the gall maker and Diomorus aiolomorphi Kamijo (Torymidae) was its inquiline.Homoporusjaponicus Ashmead (Pteromalidae) and Eupelmus sp. (Eupelmidae) are likely to be primary parasitoids of the larva ofA. rhopaloides. Eurytoma sp. (Eurytomidae),Sycophila sp. (Eurytomidae) and Norbanus sp. (Pteromalidae) are thought to be facultative hyperparasitoids. Cecidomyiidae sp. is thought to be the inquiline ofA. rhopaloides galls.Leptacis sp. (Platygastridae) probably parasitizes the larvae of Cecidomyiidae sp. Larvae of A. rhopaloides appeared in galls in July with the percentage of larvae decreasing in September, before overwintering as pupae. The growth ofD. aiolomorphi larvae within galls may be faster than that ofA. rhopaloides. The percentage of parasitoids in galls was low in July, but increased until winter. Aiolomorphus rhopaloides and D. aiolomorphi emerged from mid-April to early May, and from late April to early June, respectively. From overwintering galls, six other hymenopteran species emerged between late May and late June; one dipteran Cecidomyiidae sp. emerged between mid-April and early May.     

Opposing survivorship and fecundity effects of host phenology on the gall-forming aphid Hormaphis hamamelidis

Abstract  1. This study examined the synchronisation and roles of host and insect phenology for fundatrices of the gall-forming aphid Hormaphis hamamelidis , which forms galls on the leaves of the understorey tree Hamamelis virginiana .
2. Host and insect phenologies varied widely among years and individual host trees.
3. Most winter eggs (≈ 85%) hatched before budbreak and galls could be initiated but in 2 years there was no inherent risk for earlier hatching fundatrices in terms of greater mortality. Egg hatch was not synchronised tightly with host trees but instead seemed to occur well in advance of budbreak as a conservative, bet-hedging strategy to ensure the opportunity to form galls.
4. Fundatrices that formed galls later had higher fecundity, contrary to typical patterns for insects that feed on the first flush of leaves in temperate forests, where later herbivores have lower fecundity. Also, unlike many other systems, trees that had later budbreak had greater densities of galls.     

Distinguishing intrapopulational categories of plants by their insect faunas: galls on rabbitbrush

Within a population of rubber rabbitbrush, Chrysothamnus nauseosus, the subspecies C. nauseosus consimilis and C. nauseosus hololeucus, and a third unidentified group were better segregated by their insect galls, than by differences in plant morphology. This level of segregation was further increased when morphological measurements and counts of insect galls were analyzed simultaneously. We interpret this result to mean that plant morphology and insect distributions reflect two different, perhaps overlapping, portions of the host's genome. By using both sets of characters concurrently, rather than either set independently, we increased the portion of the plant's genome being sampled and increased the probability of detecting differences among host genotypes. Hence, knowledge of the distributions of insect galls may be useful for augmenting the level of separation, obtained by morphological measurements, among intrapopulational categories of plant genotypes. This application may be of greatest benefit when hybridization blurs morphological distinctions among plant taxa, when morphological traits are highly variable within genotypes, or when ephemeral morphological traits (e.g., leaves, flowers) are not available for measurements.     

一种花瘿蚊(双翅目:瘿蚊科)与其寄主植物对萼猕猴桃Actinidia valvata的相互关系(英文)

猕猴桃属植物Actinidia spp,自然分布于中国的亚热带地区。对萼猕猴桃A.valvata的花芽受Pseudasphond ylia 瘿蚊属一未知种的寄生而形成花芽虫瘿,这种花芽瘿由于近年来在制药工业上的应用而受到重视。在中国中南地区研究对萼猕猴桃—瘿蚊的相互关系过程中,我们的记载表明该造瘿昆虫在一定情形下可能改变其寄主植物的雌雄异株之生殖模式。野外调查与实验证明寄主植物受寄生而形成虫瘿的比例很高。但虫瘿密度却于不同沟谷间,或同一沟谷内不同植株间有异。在二条沟谷内,92％和75％植株分别被寄生,而在第3条沟谷内没有植株受寄生。受寄生的雄性植株只产生虫瘿,而受寄生的雌性植株则产生正常果与虫瘿。有迹象显示当寄主植株有虫瘿形成时其正常果数量也更多。作者认为这可能是昆虫诱导功能上雌雄异株植物形成雄性异株雌雄同株之生殖模式的少数例子     

Insect herbivores as potential causes of mortality and adaptation in gallforming insects

Summary Recent studies have suggested that plant galls benefit only the insects living in them and not the host plants, and that galls are induced by insects primarily to improve the plant as a microenvironment or a food source. The potential advantage to insects of protection from their predators and parasitoids has been considered unclear and perhaps minor in importance. However, the potential threat to gallforming insects from other insect herbivores has usually been relatively neglected. This paper notes literature and observations which suggest that herbivores may either consume or be deterred by galls. Even soft leaf galls produced by Hormaphis and Phylloxera aphids appeared to deter some herbivores in the field. The threat of herbivory to galls might help explain general patterns of gall ecology and morphology, and deserves closer attention.