昆虫的黑化机理

昆虫的黑化是昆虫出现的一种体色变异。文章介绍工业污染、激素和温度等对昆虫黑化的影响、昆虫色素形成中多巴、多巴胺、N-β丙酰多巴胺(NBAD)和N-乙酰多巴胺(NADA)等色素前体的代谢、昆虫的黑化突变以及昆虫黑化的遗传。     

水稻害虫化学生态调控研究进展

自然界中,在昆虫和植物种内与种间都存在着复杂的化学联系。开发利用生态系统中这些调控生物种内种间关系的生态功能分子,可望有效降低害虫的种群密度,从而减少化学农药的使用。本文根据目前国内外在利用生态功能分子调控水稻害虫方面的最新研究成果,分别就利用昆虫性信息素、水稻挥发物、非寄主植物提取物、化学激发子以及遗传改良水稻品种等在调控水稻害虫及其天敌中作用的研究进展进行了综述,并提出了今后的研究重点与方向。希望通过本文能促进化学生态调控技术在水稻害虫治理中的应用,以减轻水稻害虫治理对化学农药的依赖。     

昆虫迁飞的调控基础及展望

昆虫迁飞是在长期适应多变的环境过程中进化形成的一种行为对策,也是昆虫的种类和数量繁多,以及迁飞害虫经常暴发成灾的主要原因.昆虫迁飞行为的发生不仅受到外界环境因素的影响,而且受到本身生理因素的调控.目前,国内外对此类研究主要集中在生态环境、生理因素、行为学以及种群遗传学方面的调控机制.随着分子生物学技术的发展,昆虫迁飞行为发生的分子调控机制也越来越受到重视.在对国内外主要昆虫迁飞调控机制概述的基础上,对新的分子生物学技术在昆虫迁飞调控中的应用进行了探讨与展望.     

昆虫天然免疫相关基因研究进展

在长期进化过程中,昆虫形成了强大的天然免疫防御系统,即体液免疫和细胞免疫。体液免疫主要包括Toll、IMD和JAK/STAT 3条信号通路,通过信号转导及免疫途径调控免疫相关基因的表达,诱导产生抗菌肽和其他效应分子。细胞免疫由血细胞介导,主要完成对病原物的包裹、吞噬和集结等。近年来,昆虫基因组学快速发展,通过生物信息学等方法从昆虫基因组数据中已鉴定到大量免疫相关基因,对这些基因的研究加深了人们对昆虫天然免疫分子机制的认识和理解。根据基因功能,免疫相关基因分为识别、信号转导、调制器、效应分子、黑化反应、RNA干扰和其他基因等7类,这些基因通过互作来调控体液免疫和细胞免疫。本文对昆虫免疫相关基因的分类、功能及家族进化等方面的研究成果进行总结,并对今后昆虫免疫的研究重点进行了展望,以期为昆虫免疫分子机制的研究及开发新的害虫防治策略提供依据。     

黑化粘虫的形态特征及其遗传模式

本文首次研究报道了黑化粘虫Mythimna separata的形态特征及其遗传模式。所获的结果表明,黑化型与正常型粘虫的外部形态特征差异主要表现在成虫期。与正常型粘虫相比,黑化粘虫的成虫体色几乎全部变黑,正常型成虫所具有的黑色特征在黑化粘虫中已完全看不见,其他的颜色特征也由于黑化而产生了变异。卵、幼虫和蛹的颜色与正常型的没有明显的区别,只是黑化型的蛹在将羽化时的体色比正常型的深。黑化粘虫的遗传遵循孟德尔的隐性遗传规律,由位于常染色体上的单基因控制。这些结果表明,黑化型与正常型粘虫虽然仍属于同一个种,但黑化粘虫已经演变成了不同的基因型。最后,对黑化粘虫产生的原因及其在进化上的适应意义进行了讨论。     

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。     

昆虫种群的遗传调控

昆虫种群的遗传调控是利用昆虫自身生长发育的关键基因,采用性别控制开关,通过遗传转化使雄虫成为携带导致后代雌虫发育异常或雌性不育的遗传控制复合体（性别开关元件和靶标基因的复合体）．昆虫种群遗传调控是一种基于不育技术的昆虫种群控制系统,具有种类特异、环境友好和便捷高效等特点．目前为止,已经由早期的通过辐射不育方法发展到释放携带显性致死基因昆虫的方法,并在多种昆虫中获得成功．本文综述了昆虫种群遗传调控的发展历程,介绍了昆虫种群遗传调控相关的理论与方法,包括特异的调控元件、致死或缺陷基因和遗传转化体系的应用,并列举了几种昆虫种群遗传调控的实例,最后对于昆虫种群遗传调控系统中存在的问题以及可能的发展方向进行了展望．     

昆虫免疫防御分子Hemolin的研究进展

Hemolin是昆虫所特有的类免疫球蛋白,在昆虫的免疫防御通路中具有重要的作用.它本身无直接的抗菌活性,但可以延缓昆虫发病的时间.同时也可调控昆虫体内酚氧化物酶的活性,影响黑化进程.此外,Hemolin影响昆虫的发育,并且该蛋白的表达还受到昆虫体内激素的调控.综述了Hemolin各类性质和功能机制的研究进展,在此基础上,对Hemolin的功能进行探讨和总结,并对其研究前景及可能出现的问题进行分析.     

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。     

昆虫体色分化研究进展

昆虫体色分化现象在自然界中普遍存在,它给昆虫的分类等工作带来了不便。文章介绍了寄主专化论、环境因素控制论、基因控制论等有关昆虫体色分化机理的3个不同观点,着重从染色体和基因方面介绍了昆虫体色分化的研究,以及分子标记技术在其研究中的应用,并对昆虫体色分化的研究方向进行了展望。     

Genetic basis of stage-specific melanism: a putative role for a cysteine sulfinic acid decarboxylase in insect pigmentation

Melanism, the overall darkening of the body, is a widespread form of animal adaptation to particular environments, and includes bookcase examples of evolution by natural selection, such as industrial melanism in the peppered moth. The major components of the melanin biosynthesis pathway have been characterized in model insects, but little is known about the genetic basis of life-stage specific melanism such as cases described in some lepidopteran species. Here, we investigate two melanic mutations of Bicyclus anynana butterflies, called Chocolate and melanine, that exclusively affect pigmentation of the larval and adult stages, respectively. Our analysis of Mendelian segregation patterns reveals that the larval and adult melanic phenotypes are due to alleles at different, independently segregating loci. Our linkage mapping analysis excludes the pigmentation candidate gene black as the melanine locus, and implicates a gene encoding a putative pyridoxal phosphate-dependant cysteine sulfinic acid decarboxylase as the Chocolate locus. We show variation in coding sequence and in expression levels for this candidate larval melanism locus. This is the first study that suggests a biological function for this gene in insects. Our findings open up exciting opportunities to study the role of this locus in the evolution of adaptive variation in pigmentation, and the uncoupling of regulation of pigment biosynthesis across developmental stages with different ecologies and pressures on body coloration.     

Melanism and disease resistance in insects

There is growing evidence that insects in high-density populations invest relatively more in pathogen resistance than those in low-density populations (i.e. density-dependent prophylaxis). Such increases in resistance are often accompanied by cuticular melanism, which is characteristic of the high-density form of many phase polyphenic insects. Both melanism and pathogen resistance involve the prophenoloxidase enzyme system. In this paper the link between resistance, melanism and phenoloxidase activity is examined in Spodoptera larvae. In S. exempta , cuticular melanism was positively correlated with phenoloxidase activity in the cuticle, haemolymph and midgut. Melanic S. exempta larvae were found to melanize a greater proportion of eggs of the ectoparasitoid Euplectrus laphygmae than non-melanic larvae, and melanic S. littoralis were more resistant to the entomopathogenic fungus Beauveria bassiana (in S. exempta the association between melanism and fungal resistance was non-signficant). These results strengthen the link between melanism and disease resistance and implicate the involvement of phenoloxidase.     

Selection for cuticular melanism reveals immune function and life-history trade-offs in Spodoptera littoralis

Several insect species show an increase in cuticular melanism in response to high densities. In some species, there is evidence that this melanism is correlated with an up-regulation of certain immune system components, particularly phenoloxidase (PO) activity, and with the down-regulation of lysozyme activity, suggesting a trade-off between the two traits. As melanism has a genetic component, we selected both melanic and nonmelanic lines of the phase-polyphenic lepidopteran, Spodoptera littoralis, in order to test for a causative genetic link between melanism, PO activity and lysozyme activity, and to establish if there are any life-history costs associated with the melanic response. We found that, in fact, melanic lines had lower PO activity and higher lysozyme activity than nonmelanic lines, confirming a genetic trade-off between the two immune responses, but also indicating a genetic trade-off between melanism and PO activity. In addition, we found that lines with high PO activity had slower development rates suggesting that investment in PO, rather than in melanism, is costly.     

The costs of colour: plasticity of melanin pigmentation in an outbreaking polymorphic forest moth

Outside the context of industrial melanism, little is known about the physiological and ecological importance of genetic melanic polymorphisms in moths. Melanin pigments are synthesized from amino acid precursors and should therefore be costly to produce in nitrogen‐limited insects. A genetic melanic polymorphism is present in adult Malacosoma disstria Hübner (Lepidoptera: Lasiocampidae), a widespread forest moth with outbreaking population dynamics. We test the hypotheses that melanin‐based colouration is physiologically costly in M. disstria, that expression of melanin‐based colouration is a plastic trait which varies with population density and nutrition, and that the genetically based melanic phenotype is disadvantaged under nutritionally poor conditions. Two experiments were used to test these hypotheses. A field study compared pigmentation and phenotypic frequencies in moths collected from high‐ and low‐density populations. A laboratory experiment investigated the effects of larval nitrogen availability on adult pigmentation and phenotypic frequencies. High population density and nitrogen limitation reduced pigmentation and size of all moths, but phenotypic frequencies were not affected in either experiment. The effects of diet on both pigmentation and size were stronger for melanic moths than for typical moths. Our results show that adult melanism in M. disstria is physiologically costly, that colour expression is plastic despite its genetic component, and that the melanic phenotype may be disadvantaged under poor conditions but favoured under good conditions. We suggest that temporal variation in selection and trait plasticity help maintain polymorphism stability.     

Rethinking the thermal melanism hypothesis: rearing temperature and coloration in pygmy grasshoppers

Selection for efficient conversion of solar radiation to body heat has favored the evolution of dark coloration in many ectotherms. The thermal melanism hypothesis posits that dark coloration is beneficial under conditions of low ambient temperatures because it results in faster heating rates and higher body temperatures. Fast heating rates, however, may come at a cost of overheating unless compensated for by thermal physiology or behaviour. Pygmy grasshopper (Orthoptera, Tetrigidae) populations that inhabit fire-ravaged areas characterized by blackened backgrounds and hot surface temperatures due to high absorbance of solar radiation show an increased frequency of black phenotypes. I raised the progeny of wild-captured Tetrix undulata in cold and hot temperatures and used data on color patterns and survival in a greenhouse to examine whether a cold thermal environment triggered the development of melanic coloration or differently affected survival of melanic versus non-melanic individuals. My results indicate that melanism was not influenced by rearing temperature but by genes or epigenetic maternal effects. Temperature also did not affect survival. However, melanic individuals produced by melanic mothers survived longer than melanic individuals produced by non- melanic mothers, whereas non-melanic individuals produced by non-black mothers survived longer than melanic individuals produced by non-black mothers. This suggests a mismatch between color and physiology in offspring belonging to a different color morph than their mother. Future investigations into the evolution of melanism should consider conflicting selection pressures on thermal capacity and camouflage as well as the influence of correlated responses to selection on traits associated with coloration.     

Immune defence,a possible nonvisual selective factor behind the industrial melanism of moths (Lepidoptera)

It is generally believed that industrial melanism in Lepidoptera is mainly caused by differential predation by birds. In polluted areas, melanic individuals are favoured by natural selection because they are better camouflaged than pale moths on lichen‐free and sooty tree trunks. In this article, we show that, in the black arches moth (Lymantria monacha), melanic morphs have a stronger encapsulation response than pale morphs against nylon monofilament implants. This indicates that the melanic and pale morphs differ in the strength of their immune defence. The same chemical precursors and their end product, melanin pigment, are involved in the encapsulation response and in the external coloration. Thus, it seems that there may be two possible, not mutually exclusive, explanations for the frequency changes observed in the industrial melanism of moths. The dominant gene causes an increase in the amount of melanin pigment and its precursors. This increase causes two changes: an intensified immune defence as a form of improved encapsulation ability of foreign objects, and the well‐known protective dark coloration (a case of relational pleiotropy). It seems possible that industrial melanism is a by‐product of selection on the strength of immunity. In the field, these pleiotropic aspects are exceedingly difficult to distinguish from each other, and the factors may even be compensatory. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 831–838.     

More than a colour change: insect melanism,disease resistance and fecundity

A ‘dark morph’ melanic strain of the greater wax moth, Galleria mellonella, was studied for its atypical, heightened resistance to infection with the entomopathogenic fungus, Beauveria bassiana. We show that these insects exhibit multiple intraspecific immunity and physiological traits that distinguish them from a non-melanic, fungus-susceptible morph. The melanic and non-melanic morphs were geographical variants that had evolved different, independent defence strategies. Melanic morphs exhibit a thickened cuticle, higher basal expression of immunity- and stress-management-related genes, higher numbers of circulating haemocytes, upregulated cuticle phenoloxidase (PO) activity concomitant with conidial invasion, and an enhanced capacity to encapsulate fungal particles. These insects prioritize specific augmentations to those frontline defences that are most likely to encounter invading pathogens or to sustain damage. Other immune responses that target late-stage infection, such as haemolymph lysozyme and PO activities, do not contribute to fungal tolerance. The net effect is increased larval survival times, retarded cuticular fungal penetration and a lower propensity to develop haemolymph infections when challenged naturally (topically) and by injection. In the absence of fungal infection, however, the heavy defence investments made by melanic insects result in a lower biomass, decreased longevity and lower fecundity in comparison with their non-melanic counterparts. Although melanism is clearly correlated with increased fungal resistance, the costly mechanisms enabling this protective trait constitute more than just a colour change.     

The Genetic,Morphological, and Physiological Characterization of a Dark Larval Cuticle Mutation in the Butterfly,Bicyclus anynana

Studies on insect melanism have greatly contributed to our understanding of natural selection and the ultimate factors influencing the evolution of darkly pigmented phenotypes. Research on several species of melanic lepidopteran larvae have found that low levels of circulating juvenile hormone (JH) titers are associated with a melanic phenotype, suggesting that genetic changes in the JH biosynthetic pathway give rise to increased deposition of melanin granules in the cuticle in this group. But does melanism arise through different molecular mechanisms in different species? The present study reports on a Bicyclus anynana (Lepidoptera: Nymphalidae) dark larvae single locus mutation, in which larvae exhibit a darker cuticle relative to wild type. Unlike other lepidopteran melanic larvae mutations, this one is autosomal recessive and does not appear to involve a deficiency in JH titers. Unlike JH deficiency mutants, dark larvae mutants display similar growth rates and sexual behaviors as wild type, and topical application of a JH analogue failed to rescue the wild type cuticular coloration. Finally, transmission electron microscopy showed that sclerotization or deposition of diffuse melanin, rather than deposition of melanin granules, produces the dark coloration found in the cuticle of this species. We conclude that different molecular mechanisms underlie larval melanism in different species of Lepidoptera.     

Melanic mutation causes a fitness decline in bean beetles infected by Wolbachia

Wolbachia cannot live outside a host, which is thought to be the reason for host‐Wolbachia coevolution toward benign parasitism, especially because the fitness of Wolbachia is traded against its host's fitness. Insect melanism has been reported to have a positive effect on pathogen resistance, but melanic mutants of Callosobruchus analis (Fabricius) and Callosobruchus chinensis (L.) (Coleoptera: Chrysomelidae) are infected with Wolbachia. Callosobruchus chinensis is infected with CI‐inducing Wolbachia, and melanic mutants exhibit fitness decline. Interestingly, this decline is not observed in C. analis melanic mutants that are infected with CI‐free Wolbachia. Our research question is whether the infection of CI‐inducing Wolbachia causes fitness decline of melanic hosts in C. analis. We examined fecundity, fertility, and longevity of C. analis melanic mutants and compared them between uninfected and infected hosts with CI‐inducing Wolbachia. Infected melanic mutants of C. analis exhibited fitness decline leading to reduced hatch rates even when parental combinations were compatible. Wolbachia can invade a host population by causing CI to decrease the fraction of uninfected hosts, but melanic mutant hosts decrease the number of infected hosts through fitness decline. Nevertheless, the melanism in hosts is not able to stop Wolbachia invasion in C. analis.     

A proposed selective mechanism based on metal chelation in industrial melanic moths

Industrial melanism, a phenomenon observed in some moths and especially in the case of the peppered moth (Biston betularia), has received much attention as an example of Darwinian evolution in action. The rapid rise in the proportion of the darker melanic form of the adult moth coincided with the advent of atmospheric pollution resulting from industrialization, and was ascribed to the improved camouflage of the melanotic insects against a background blackened by soot, which conferred a selective advantage in the avoidance of predation by birds. The topic of the increase in melanization during the initial period of industrial expansion and the reversal of the process after the introduction of the Clean Air Act has received much attention. Although there is sound experimental evidence to support selective avian predation as a major mechanism to account for the changes in the relative frequency of melanics, it is not clear that this is the only selective factor involved in industrial melanism. It is possible that other processes may have made a contribution to the preponderance of melanic variants. In the present study, the hypothesis is advanced that melanization may have conferred a selective advantage by protecting the insects from the toxic effects of metals by virtue of the strong metal chelating action of melanin. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 298–301.