植物病媒昆虫的翅型分化

翅多型现象是昆虫非遗传多型性的一种表现,包括不具飞行能力的短翅型或无翅型,以及可以进行长距离迁飞的长翅型或有翅型。翅多型现象常发生在可以携带病原并将其传播给植物宿主的媒介昆虫中,对植物病害的时空分布与暴发有重要影响。本文从翅型分化的遗传规律、诱导因素、分子机制和伴随翅型分化的其他生理表现4个方面,对植物病原主要传播媒介蚜虫和飞虱的翅型分化研究进行综述和梳理。昆虫翅型分化的诱导因素主要包括温度、湿度和光周期等非生物因素以及虫口密度、宿主营养、病毒等生物因素;而其内在的分子机制大多是通过胰岛素/胰岛素样生长因子信号(IIS)通路、c-Jun氨基末端激酶(c-Jun NH 2-terminal kinase,JNK)信号通路、Wingless和嗅觉受体SaveOrco等调控。翅型分化的同时伴随着生理状态的变化,表现为短翅型具有更强的繁殖能力和长翅型含有更丰富的飞行肌结构成分。目前,昆虫翅型分化的研究尚不够完善,有许多需要解答的问题,如找到胰岛素/胰岛素样生长因子信号通路中真正发挥功能的靶基因,JNK如何调控翅型分化以及虫媒病毒影响媒介昆虫翅型的分子机理。本综述可为控制虫媒病原的传播以及其他昆虫翅多型的研究提供参考。     

昆虫翅型分化的调控及翅多型性的进化

翅多型现象普遍存在于各昆虫类群,一些学者就昆虫翅多型进行了大量的研究工作。根据昆虫翅型的分化,可划分为长翅型和短翅型,长翅型具飞行能力,而短翅型不能飞行。一些昆虫种类,如蚜虫,出现无翅个体,被称为无翅型。除飞行能力外,长翅型和短翅型在行为、生理等方面也存在差异。文章主要就环境因素对翅型分化的影响、翅多型的内分泌控制机理、翅多型的遗传机制及其进化等作一概述。     

昆虫非遗传多型性研究进展

非遗传多型性是指同一基因型或同一基因组通过外界环境诱导可产生两种或者多种不连续表型的现象。该现象在昆虫中已有报道,如变态、季节性非遗传多型、社会性昆虫的等级制等。昆虫通过非遗传多型性做出应答,通过表型改变来适应环境并利用周围环境物质以达到躲避天敌从而进行生存繁衍的目的。因此,非遗传多型性是昆虫种类繁多、数量庞大的主要因素之一。近几十年来,非遗传多型性日益受到广泛关注,从最初对现象的描述,到诱导该现象产生的可能因素的实验验证,至目前大数据时代下利用二代测序技术、基因敲除和RNA干扰等技术揭示其分子机制。本文对近年来非遗传多型性在昆虫中的研究进展进行了总结,并对未来的趋势进行了展望。     

沙蟋翅多型性的调控机理

沙蟋Gryllus firmus Zera&Denno成虫的后翅有长翅和短翅2种类型,是翅多型性机理研究的极佳模式昆虫。长翅成虫从第5日龄开始迁飞,而短翅成虫的主要特点是繁殖。除了翅的表型差异外,长翅成虫的飞行肌发达,呈褐色;卵巢幼小,直到飞行停止后(大约在10d以后)才开始发育。而短翅成虫的飞行肌退化并呈乳白色;卵巢在第4日龄就发育成熟,表现为卵巢硕大。对翅多型性机理的深入研究,将有利于了解沙蟋迁飞和扩散的内在机理,为准确地预测预报该虫的发生提供重要的理论和实际依据。文章概述沙蟋翅多型性与外界环境的相互关系,以及体内生化代谢和内分泌激素等的变化对该虫迁飞和生殖的影响和作用,进而探讨翅多型的遗传机制和进化意义等问题。     

社会性昆虫级型和行为分化机制研究进展

社会性的出现是生物演化过程中的重要革新, 理解社会性的演化和调控机制具有重要的理论和实际意义。社会性昆虫的个体间有着明显的级型分化和劳动分工, 这有利于它们适应复杂的环境变化。理解社会性昆虫如何产生不同的形态、行为和生活史特性, 一直是进化和发育生物学的重要目标。随着测序技术的不断更新及生物信息学的快速发展, 已经有众多关于社会性昆虫级型和行为分化机制的研究报道。本文通过整理社会性昆虫研究的已有成果, 从环境因素、生理调控和分子机制等方面对社会性昆虫级型和行为分化机制相关研究进展进行了综述, 并对未来的研究方向做出了展望。根据现有证据, 社会性昆虫所生活的生物环境(食物营养、信息素、表皮碳氢化合物)和非生物环境(温度、气候等)均能直接或间接影响社会性昆虫级型和行为的分化; 保幼激素、蜕皮激素、类胰岛素及生物胺等内分泌激素和神经激素对社会性昆虫的级型和行为分化也有重要的调控作用; 此外, 遗传因素、新基因等DNA序列或基因组结构上的变化以及表观遗传修饰、基因的差异表达等基因调控机制均能不同程度地影响社会性昆虫的行为分化。本文建议加强昆虫纲其他社会性类群如半翅目蚜虫和缨翅目蓟马等的社会性行为及其演化机制的研究, 以加深对社会性昆虫起源及其行为演化的理解和认识。     

昆虫翅多型的分子调控机制研究进展

生物如何适应复杂的环境变化是生物学研究的重要科学问题。其中,昆虫在长期的进化过程中形成了翅多型现象以适应复杂的环境,由此通过调控翅型的转换,对不同的环境条件做出响应,从而优化资源分配以平衡迁移或繁殖的需求,但目前对其分子调控机制仍不十分清楚。本文基于国内外最新研究进展,结合作者自己的研究,系统地综述了昆虫体内参与翅型转换调控的多种途径的研究进展,包括胰岛素信号通路、蜕皮激素信号通路、保幼激素信号通路、JNK信号通路、生物胺信号通路和病毒基因水平转移,指出了未来发展的方向。这些研究成果不仅对于进一步阐明昆虫翅多型的分子机制具有指导意义,也为开发以翅型调控为主要内容的害虫综合治理技术提供参考。     

昆虫体色多型的分子调控机制

体色多型普遍存在于各昆虫类群,体色多型不仅是生物多样性的体现,而且多样的着色模式对于昆虫本身具有重要的生物学意义.研究体色多型对探讨昆虫遗传多态性、适应机制及生物进化等具有重要的意义.本文主要从昆虫体色多型的分子调控机制进行综述,以期为今后探讨昆虫多态性、适应机制及生物进化提供参考.     

温度对桃蚜和萝卜蚜翅型分化的影响

采用恒温试验,自然变温试验和大田系绕调查相结合的研究方法,探讨了温度对桃蚜和萝卜蚜孤雌胎生型翅型分化的影响,结果表明低温有助于翅的发育,高温则对翅的发育有抑制作用,但低温对翅发育的促进作用在桃蚜中比在萝卜蚜中要强得多。在桃蚜中还证实母蚜体内的仔蚜胚胎期及仔蚜都可感受温度的作用从而对仔蚜的翅型分化产生影响。根据本文结果并综合文献中有关报道,作者认为在确定蚜虫翅型分化与环境因子的关系时,温度是一个不可忽略的重要因子。     

遗传和环境因素对曲脉姬蟋亚热带种群翅型分化的影响

曲脉姬蟋Modicogryllus confirmatus Walker具有明显的翅二型现象。为探究环境及遗传如何影响曲脉姬蟋亚热带种群的翅型分化,对饲养于不同光周期、温度和密度条件下若虫羽化后的翅型比进行了调查,并对长、短翅型蟋蟀进行了3代遗传筛选和杂交试验,研究了光周期、温度、种群密度和遗传对曲脉姬蟋广西种群翅型分化的影响。结果表明:光周期和种群密度对曲脉姬蟋的翅型分化均无影响,而温度对其翅型分化具有调控作用。正常范围内的温度变化(25℃、30℃)对其翅型分化无显著影响,而35℃的极高温则显著降低曲脉姬蟋的长翅率,说明其翅型分化并不是对季节变化的适应,而高温胁迫可引起短翅化。对不同翅型进行了3代筛选,结果表明,往短翅型选拔会引起雌、雄虫的短翅率都明显下降,而往长翅型选拔时,雌、雄虫的短翅率均维持在极低水平;不同亲本组合的后代间的长翅率有差异,说明曲脉姬蟋的翅型分化可能受多基因调控。     

蜜蜂级型分化机理

蜜蜂Apis spp.能有效地为多种植物及农作物授粉, 具有重要的经济和生态价值; 蜜蜂作为高度真社会性昆虫, 已成为社会生物学研究的模式生物。社会性昆虫的生殖劳动分工具有重要的进化意义, 而级型分化是形成生殖劳动分工的基础。近年来, 关于蜜蜂级型分化的研究已取得诸多重要成果, 其机理也得到了较为深入的阐释。营养差异引发蜜蜂幼虫的级型分化。蜂王浆中的主要蛋白组分之一--Royalactin是诱导蜂王发育的关键营养因子, 而脂肪体细胞的表皮生长因子受体介导了Royalactin的这种蜂王诱导作用。DNA甲基化是重要的表观遗传机制之一, 且与个体发育和疾病发生紧密相关, 近来的研究表明DNA甲基化在蜜蜂级型分化过程中发挥重要的调控作用。此外, 越来越多的研究进一步深化了人们对内分泌系统调节级型分化作用的认识。本文从关键营养因子调控、 表观遗传调控和内分泌调节3方面综述蜜蜂级型分化的机理, 并对未来的研究提出可能的方向。     

A sex-linked locus controls wing polymorphism in males of the pea aphid,Acyrthosiphon pisum (Harris)

Discrete variation in wing morphology is a very common phenomenon in insects and has been used extensively in the past 50 years as a model to study the ecology and evolution of dispersal. Wing morph determination can be purely genetic, purely environmental, or some combination of the two. The precise genetic determinants of genetically based wing morph variation are unknown. Here we explore the genetic basis of wing polymorphism in the pea aphid, which can produce either winged or wingless males. We confirm that three types of pea aphid clones coexist in natural populations, those producing winged males only, those producing wingless males only, and those producing a mixture of both. A Mendelian genetic analysis reveals that male wing polymorphism in pea aphids is determined by a single locus, two alleles system. Using microsatellite loci of known location, we show that this locus is on the X chromosome. The existence of a simple genetic determinism for wing polymorphism in a system in which genetic investigation is possible may help investigations on the physiological and molecular mechanisms of genetically-based wing morph variation. This locus could also be used in the search for genes involved in the wing polyphenism described in parthenogenetic females and to investigate the interplay between polymorphisms and polyphenisms.     

Male-specific flight apparatus development in Acyrthosiphon pisum (Aphididae, Hemiptera, Insecta): comparison with female wing polyphenism

Wing polymorphisms observed in many Insecta are important topics in developmental biology and ecology; these polymorphisms are a consequence of trade-offs between flight and other abilities. The pea aphid, Acyrthosiphon pisum, possesses 2 types of wing polymorphisms: One is a genetic wing polymorphism occurring in males, and the other is an environmental wing polyphenism seen in viviparous females. Although genetic and environmental cues for the 2 wing polymorphisms have been studied, differences in their developmental regulation have not been elucidated. In particular, there is little knowledge regarding the developmental processes in male wing polymorphism. Therefore, in this study, the development of flight apparatuses and external morphologies was compared among 3 male wing morphs (winged, wingless, and intermediate). These male developmental processes were subsequently compared with those of female wing morphs. Developmental differences between the male and female polymorphisms were identified in flight muscle development and degeneration but not in wing bud development. Furthermore, the nymphal periods of wingless and intermediate males were significantly shorter than that of winged males, indicating the adaptive significance of male winglessness. Overall, this study indicates that the male and female wing polymorphisms are based on different regulatory systems for flight apparatus development, which are probably the result of different adaptations under different selection pressures.     

Genetic variation for an aphid wing polyphenism is genetically linked to a naturally occurring wing polymorphism

Many polyphenisms are examples of adaptive phenotypic plasticity where a single genotype produces distinct phenotypes in response to environmental cues. Such alternative phenotypes occur as winged and wingless parthenogenetic females in the pea aphid (Acyrthosiphon pisum). However, the proportion of winged females produced in response to a given environmental cue varies between clonal genotypes. Winged and wingless phenotypes also occur in males of the sexual generation. In contrast to parthenogenetic females, wing production in males is environmentally insensitive and controlled by the sex-linked, biallelic locus, aphicarus (api). Hence, environmental or genetic cues induce development of winged and wingless phenotypes at different stages of the pea aphid life cycle. We have tested whether allelic variation at the api locus explains genetic variation in the propensity to produce winged females. We assayed clones from an F2 cross that were heterozygous or homozygous for alternative api alleles for their propensity to produce winged offspring. We found that clones with different api genotypes differed in their propensity to produce winged offspring. The results indicate genetic linkage of factors controlling the female wing polyphenism and male wing polymorphism. This finding is consistent with the hypothesis that genotype by environment interaction at the api locus explains genetic variation in the environmentally cued wing polyphenism.     

Parasitoids induce production of the dispersal morph of the pea aphid, Acyrthosiphon pisum

In animals, inducible morphological defences against natural enemies mostly involve structures that are protective or make the individual invulnerable to future attack. In the majority of such examples, predators are the selecting agent while examples involving parasites are much less common. Aphids produce a winged dispersal morph under adverse conditions, such as crowding or poor plant quality. It has recently been demonstrated that pea aphids, Acyrthosiphon pisum, also produce winged offspring when exposed to predatory ladybirds, the first example of an enemy‐induced morphological change facilitating dispersal. We examined the response of A. pisum to another important natural enemy, the parasitoid Aphidius ervi, in two sets of experiments. In the first set of experiments, two aphid clones both produced the highest proportion of winged offspring when exposed as colonies on plants to parasitoid females. In all cases, aphids exposed to male parasitoids produced a higher mean proportion of winged offspring than controls, but not significantly so. Aphid disturbance by parasitoids was greatest in female treatments, much less in male treatments and least in controls, tending to match the pattern of winged offspring production. In a second set of experiments, directly parasitised aphids produced no greater proportion of winged offspring than unparasitised controls, thus being parasitised itself is not used by aphids for induction of the winged morph. The induction of wing development by parasitoids shows that host defences against parasites may also include an increased rate of dispersal away from infected habitats. While previous work has shown that parasitism suppresses wing development in parasitised individuals, our experiments are the first to demonstrate a more indirect influence of parasites on insect polyphenism. Because predators and parasites differ fundamentally in a variety of attributes, our finding suggests that the wing production in response to natural enemies is of general occurrence in A. pisum and, perhaps, in other aphids.     

Relative contribution of genetic and environmental factors to determination of wing morphs of the brown planthopper Nilaparvata lugens

Wing dimorphism is a fascinating feature of the ability of insects to adapt to environments. The brown planthopper (BPH) Nilaparvata lugens, a serious pest of rice, can switch between the long- and short-winged morphs. It has been known that environmental factors can affect the wing morph of BPH. However, it is still unclear whether the effect of environment is dependent on BPH genetic backgrounds or not. In the present study, we established the pure-bred lineages of short- and long-winged BPHs via multigenerational selection, and we found that survival and fecundity were similar between these 2 lineages. Wing morphs of the pure-bred lineages were almost fully dependent on genetics, but independent of the environmental factors, nymphal density and rice plant stage, 2 key factors affecting BPH wing morphs. In the unselected BPH population, short- and long-winged morphs were produced depending on those 2 environmental factors, indicating the contribution of environment to wing morph. In the wing-selected lineages, 4 developmental regulated genes of wing, NlInR1, NlInR2, NlAkt, and NlFoxo were expressed stably in the short-winged adults, but almost silenced in the long-winged adults. However, all these genes were expressed normally with a similar level in both the short- and long-winged adults in an unselected population except NlFoxo. The pure-bred lineages of long- and short-winged morphs exhibited different expression patterns of wing development-regulated genes, suggesting the genetic determination of wing morphs. Effects of environmental factors on wing morphs occurred only in the genetic mix population.     

Development of a wingless morph in the ladybird beetle, Adalia bipunctata

SUMMARY Many taxa of winged insects have independently lost the ability to fly and often possess reduced wings. Species exhibiting natural variation in wing morphology provide opportunities to investigate the genetics and developmental processes underlying the evolution of alternative wing morphs. Although many wing dimorphic species of beetles are known, the underlying mechanisms of variation are not well understood in this insect order. Here, we examine wing development of wild type and natural wingless morphs of the two-spot ladybird beetle, Adalia bipunctata . We show that both pairs of wings are distally truncated in the wingless adults. A laboratory population of the wingless morph displays heritable variation in the degree of wing truncation, reflecting reduced growth of the larval wing discs. The coexistence of variable wingless morphs supports the idea that typical monomorphic wingless insects may be the result of a gradual evolution of wing loss. Gene expression patterns in wing discs suggest that the conserved gene network controlling wing development in wild-type Adalia is disrupted in the dorsoventral patterning pathway in the wingless morphs. Previous research on several species of ant has revealed that the anteroposterior wing patterning pathway is disrupted in wingless workers. Future investigations should confirm whether interruptions in both taxa are limited to the patterning pathways found thus far, or whether there are also shared interruption points. Nevertheless, our results highlight that diverse mechanisms of development are likely to underlie the evolution of wingless insects.