中国昆虫表皮碳氢化合物与昆虫化学分类学的研究进展

基于表皮碳氢化合物的昆虫化学分类学已成为昆虫分类的一个重要工具。本文首先简述了世界范围内昆虫化学分类学的起源、发展及应用领域;其次在收集的大量文献基础上,综合分析了我国已开展昆虫表皮碳氢化合物组成研究的昆虫种类、研究内容、取样与进样技术、数据的统计分析、应用领域和存在的问题。针对我国昆虫表皮碳氢化合物研究进展及其在昆虫分类中应用的现状,提出了今后我国在昆虫化学分类领域的研究方向,并展望其应用前景。     

一种可用于桔小实蝇表皮碳氢化合物分析的固体进样技术

【目的】利用表皮碳氢化合物进行昆虫鉴定是目前昆虫分类学中非常活跃的研究方向之一。昆虫表皮碳氢化合物的液体浸提及进样技术是GC-MS常规的进样方法之一,随着现代技术的发展,固体进样也被广泛地用于昆虫表皮碳氢化合物的鉴定及昆虫的分类与进化分析。本研究探索利用固体进样法开展实蝇表皮碳氢化合物的提取与分析,探索并获得简便可行的取样与进样技术,为检疫性实蝇的快速鉴定提供新方法。【方法】以桔小实蝇Bactrocera dorsalis(Hendel)成虫为材料,采用固体进样与液体浸提进样方法,开展实蝇表皮碳氢化合物组成成分鉴定与分析;比较获得桔小实蝇成虫取样部位,并分析标本保存时间对桔小实蝇表皮碳氢化合物组成的影响。【结果】利用固体进样技术获得的桔小实蝇翅表皮碳氢化合物种类(39)较液体进样的(36)略多,且翅和中足的表皮碳氢化合物含量分别为0.18~23.37和0.03~4.13 ng/μg,极显著地高于用液体进样法检测到相同部位的化合物含量(分别为0.05~3.93和0.01~0.68 ng/μg)(P0.01),而且前者的检测灵敏度是后者的1.7~10.8倍;同时,翅样品的表皮碳氢化合物含量显著地高于足中的含量(P0.01)。利用固体进样获得的桔小实蝇成虫翅、足或胸部的表皮碳氢化合物图谱与对照处理所得图谱相似。干燥保存30 d的桔小实蝇翅的碳氢化合物的种类及含量与新鲜标本(保存0 d)无显著性差异(P0.05);但保存30,60和150 d标本的碳氢化合物存在显著差异(P0.05),主要是含量的差异。【结论】研究表明,利用固体进样检测昆虫表皮碳氢化合物的灵敏度较高,而且桔小实蝇成虫的主要表皮碳氢化合物种类较稳定,因此可以作为我国口岸检疫性实蝇表皮碳氢化合物获取与分析的一种新方法,而且检测中翅是固体进样的较合适取样部位。     

雄性昆虫性信息素的研究进展

全面概述雄性昆虫性信息素的研究进展。主要从性信息素的释放机制及其作用、已发现的雄性昆虫性信息素及其化学结构进行阐述。通过不同部位释放的雄性昆虫性信息素种类是多样的,但其化学结构的差异又是细微的,雄性昆虫则利用这些差异来引诱雌性昆虫、促进交配和生殖隔离等。最后对雄性昆虫性信息素的研究作了展望。     

我国昆虫信息素研究与应用的进展

用昆虫信息素防治害虫是20世纪60年代以来发展的一种治虫新技术。由于具有高效、无毒、没有污染、不伤益虫等优点,国内外对昆虫信息素的研究与应用都很重视。昆虫信息素（InsectPhernmones）是同种昆虫个体之间在求偶、觅食、栖息、产卵、自卫等过程中起通讯联络作用的化学信息物质,主要有性信息素（SexPheromones）、聚集信息素（AggregationPhero－mones）、示踪信息素（TrailPheromones）、报警信息素（AlarmPheromones）、疏散信息素（EpideicticPheromone）以及蜂王情息素（QueenPhero－mones）、那氏情息素（NosanovPheromon…     

昆虫的性信息素及其应用

较系统地介绍了昆虫性信息素领域取得的成就,包括昆虫性信息素的种类、化学、生物学、结构鉴定及信息应用等,并进行了展望。     

昆虫的表皮腺体

<正> 一、昆虫的表皮腺体及其分类 昆虫的表皮腺体是由昆虫体壁表皮细胞层中的某些特化的具有外分泌功能的细胞组成。这些表皮腺体除口腔中的唾腺在昆虫的消化、造丝等方面的功能之外,主要具有分泌种信息素和防御性物质的功能。近年来,由于人们逐渐认识到昆虫表皮腺体的分泌物对昆虫行为和生理的重要性,特别是利用昆虫性信息素在控制害虫和研究害虫的成功,加上扫描和透射电子显微镜使用的普及,促进了对表皮腺体的研     

生物的化学通讯

生物个体（同种或异种）相互之间的有着化学信息联系,昆虫,特别是社会性昆虫化学通讯是其沟通信息的主要方式,我们关于化学通讯的知识大多来自于对昆虫信息素的研究,近来的研究已揭示在植物和植物以及植物和动物之间也存在着化学信息交流。     

水生无脊椎动物的化学通讯

水生无脊椎动物的化学通讯邱高峰，堵南山，赖伟（华东师范大学生物学系上海２０００６２）关键词水生无脊椎动物，化学通讯，信息素，利己素，利它素陆生无脊椎动物昆虫化学通讯的研究已经取得了蓬勃发展，迄今为止已被分离、纯化和鉴定的昆虫信息物质有百余种，利用昆虫...     

水椰八角铁甲不同日龄和性别成虫表皮碳氢化合物的种类及动态

水椰八角铁甲Octodonta nipae(Maulik)是一种严重为害棕榈科植物的外来入侵害虫。为了明确日龄和性别等个体因素对水椰八角铁甲成虫表皮碳氢化合物的影响,本研究采用化学萃取法和气相色谱-质谱联用技术,分析了不同日龄的水椰八角铁甲雌、雄成虫表皮碳氢化合物的组成和含量。结果表明,水椰八角铁甲成虫的表皮碳氢化合物由21种C_(18)-C_(30)的碳氢化合物构成,包括11种正构烷烃、2种单甲基烷烃、1种多甲基烷烃、3种正构烯烃、1种单甲基烯烃和3种环烷烃。不同日龄和性别成虫的表皮碳氢化合物的组成种类相同,但是一些组分的含量有明显变化,例如1 d、5 d、10 d和15 d雌、雄成虫之间正二十四烷、正二十六烷、2-甲基十九烷和1-二十烯的含量存在显著差异。水椰八角铁甲成虫表皮碳氢化合物具有性二型,如性成熟雄虫的表皮的正二十烷、正二十一烷、正二十四烷、正二十六烷、2-甲基十九烷、10-甲基二十烷、1-二十烯和1-二十二烯的含量显著高于性成熟雌虫。本文的研究结果可为检验检疫过程中水椰八角铁甲近缘种,以及日龄、性别和性成熟的快速鉴定提供支持。     

我国昆虫信息素研究的回顾,思考和启迪

回顾昆虫信息素的研究历程,自1959年西德著名科学家A.布特南博士首次鉴定出第一个昆虫性信息素化学结构—蚕蛾醇后,至今已30年有余,无论在信息素的化学,生物学,生理学,生态学和植物保护等方面都取得了巨大成就;先后鉴定出信息素化学结构的昆虫种类达700余种;有数百种昆虫性信息素和引诱剂应用于害虫测报和防治;全世界将近有十多家公司生产和销售几十种昆虫信息素诱芯和各种诱捕器的产品,给农林害虫综合治理带来了巨大     

Natural Variation in the Strength and Direction of Male Mating Preferences for Female Pheromones in Drosophila melanogaster

Many animal species communicate using chemical signals. In Drosophila, cuticular hydrocarbons (CHCs) are involved in species and sexual identification, and have long been thought to act as stimulatory pheromones as well. However, a previous study reported that D. melanogaster males were more attracted to females that were lacking CHCs. This surprising result is consistent with several evolutionary hypotheses but is at odds with other work demonstrating that female CHCs are attractive to males. Here, we investigated natural variation in male preferences for female pheromones using transgenic flies that cannot produce CHCs. By perfuming females with CHCs and performing mate choice tests, we found that some male genotypes prefer females with pheromones, some have no apparent preference, and at least one male genotype prefers females without pheromones. This variation provides an excellent opportunity to further investigate the mechanistic causes and evolutionary implications of divergent pheromone preferences in D. melanogaster males.     

Chemical convergence between a guild of facultative myrmecophilous caterpillars and host plants

1. Ants exert strong selective pressure on herbivorous insects, although some caterpillars can live in symbiosis with them using chemical defensive strategies.
2. We investigated the adaptive resemblance of cuticular hydrocarbons (CHCs) in multitrophic systems involving a guild of facultative myrmecophilous caterpillar species (Lepidoptera: Lycaenidae), tending ants (Hymenoptera: Formicidae), and host plants from three families. We hypothesised that the CHCs of the caterpillars would resemble those of their host plants (chemical camouflage).
3. We analysed CHCs using gas chromatography/mass spectrometry. Morisita's similarity index (SI) was used to compare CHC profiles of caterpillar species with different types of ant associations (commensal or mutualistic), ants, and host plants.
4. We found strong convergence between caterpillars' CHCs and plants, especially for commensal species that do not provide secretion rewards for ants. Moreover, we found unexpected chemical convergence among mutualistic (trophobiotic) caterpillar species that offer caloric reward secretions to ants.
5. These results show that the studied caterpillars acquire CHCs through their diet and that they vary according to host plant species and type of ant association (commensalism or mutualism). This ‘chemical camouflage’ of myrmecophilous caterpillars may have arisen as a defensive strategy allowing coexistence with ants on plants, whereas ‘chemical conspicuousness’ may have evolved in the context of honest signalling between mutualistic partners.
6. We suggest the existence of chemical mimicry among myrmecophilous species, especially between mutualistic caterpillars. Cuticular chemical mixtures can play a key adaptive role in decreasing ant attacks and increasing caterpillar survival in multimodal sensory systems.

     

‘Does my Diet Affect my Perfume?’ Identification and Quantification of Cuticular Compounds in Five Drosophila melanogaster Strains Maintained over 300 Generations on Different Diets

Cuticular hydrocarbons (CHCs) in Drosophila melanogaster represent the basis of chemical communication being involved in many important biological functions. The aim of this study was to characterize chemical composition and variation of cuticular profiles in five D. melanogaster strains. These strains were reared for approximately 300 generations on five diets: standard cornmeal medium and substrates prepared with apple, banana, tomato, and carrot. Differences in quantity and/or quality in CHCs were assumed as a result of activation of different metabolic pathways involved in food digestion and adaptations to the particular diet type. In total, independently of sex and strain, 66 chemical compounds were identified. In females of all strains, 60 compounds were identified, while, in males, 47 compounds were extracted. Certain new chemical compounds for D. melanogaster were found. MANOVA confirmed that CHC amounts significantly depend on sex and substrates, as well as on their interactions. Discriminant analysis revealed that flies belonging to ‘apple’ and ‘carrot’ strains exhibited the most noticeable differences in CHC repertoires. A non‐hydrocarbon pheromone, cis‐vaccenyl acetate (cVA) also contributed to the variation in the pheromone bouquet among the strains. Variability detected in CHCs and cVA may be used in the explanation of differences in mating behaviour previously determined in analyzed fly strains.     

Wax,sex and the origin of species: Dual roles of insect cuticular hydrocarbons in adaptation and mating

Evolutionary changes in traits that affect both ecological divergence and mating signals could lead to reproductive isolation and the formation of new species. Insect cuticular hydrocarbons (CHCs) are potential examples of such dual traits. They form a waxy layer on the cuticle of the insect to maintain water balance and prevent desiccation, while also acting as signaling molecules in mate recognition and chemical communication. Because the synthesis of these hydrocarbons in insect oenocytes occurs through a common biochemical pathway, natural or sexual selection on one role may affect the other. In this review, we explore how ecological divergence in insect CHCs can lead to divergence in mating signals and reproductive isolation. We suggest that the evolution of insect CHCs may be ripe models for understanding ecological speciation.     

Intracolony chemical communication in social insects

Chemical messengers are the primary mode of intracolony communication in the majority of social insect species. Chemically transmitted information plays a major role in nestmate recognition and kin recognition. Physical and behavioral castes often differ in chemical signature, and queen effects can be significant regulators of behavior and reproduction. Chemical messengers themselves differ in molecular structure, and the effects on behavior and other variables can differ as a consequence of not only molecular structure of the chemical messenger itself but also of its temporal expression, quantity, chemical blends with other compounds, and effects of the environment. The most studied, and probably the most widespread, intracolony chemical messengers are cuticular hydrocarbons (CHCs). CHCs are diverse and have been well studied in social insects with regard to both chemical structure and their role as pheromones. CHCs and other chemical messengers can be distributed among colony members via physical contact, grooming, trophallaxis, and contact with the nesting substrate. Widespread intracolony distribution of chemical messengers gives each colony a specific odor whereby colony members are integrated into the social life of the colony and non-members of the colony are excluded. Colony odor can vary as a function of genetic diversity within the colony, and the odor of a colony can change as a function of colony age and environmental effects. Chemical messengers can disseminate information on the presence of reproductives and fertility of the queen(s) and workers, and queen pheromone can play a significant role in suppressing reproduction by other colony members. New analytical tools and new avenues of investigation can continue to expand knowledge of how individual insects function as members of a society and how the society functions as a collective.     

Cuticular hydrocarbon profiles of codling moth larvae,Cydia pomonella (Lepidoptera: Tortricidae), reflect those of their host plant species

The cuticular hydrocarbons (CHCs) of codling moth larvae collected in the field from their host plant species, apple and walnut, were analyzed and compared with the CHCs of fruits from these two phylogenetically distant hosts. The CHC profiles of the larvae consisted solely of n‐alkanes (C₂₃–C₃₁) and differed quantitatively between host populations. Amounts of the CHCs from the walnut‐collected larvae were shifted towards longer‐chain alkanes compared to those from apple‐collected larvae. A similar shift was observed for the CHC profiles of walnut and apple fruits. Analysis of the CHCs of larvae reared on artificial diet, in comparison with hydrocarbons from the diet, confirmed that larval CHCs scarcely reflect hydrocarbons from the food source. This finding indicates that the larval hydrocarbons must be biosynthesized to a large degree by the insect, rather than being gained directly from its diet. Hence, codling moth populations from apple and walnut each synthesize their own CHC profiles, which largely resemble those of their respective host plant, yielding a potential tool of chemical camouflage from certain natural antagonists of the larvae. The findings of the present study, together with recent molecular population analyses, provides evidence for a process that might ultimately lead to sympatric speciation of this herbivore species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 376–384.     

More to legs than meets the eye: Presence and function of pheromone compounds on heliothine moth legs

Chemical communication is ubiquitous in nature and chemical signals convey species-specific messages. Despite their specificity, chemical signals may not be limited to only one function. Identifying alternative functions of chemical signals is key to understanding how chemical communication systems evolve. Here, we explored alternative functions of moth sex pheromone compounds. These chemicals are generally produced in, and emitted from, dedicated sex pheromone glands, but some have recently also been found on the insects' legs. We identified and quantified the chemicals in leg extracts of the three heliothine moth species Chloridea (Heliothis) virescens, Chloridea (Heliothis) subflexa and Helicoverpa armigera, compared their chemical profiles and explored the biological function of pheromone compounds on moth legs. Identical pheromone compounds were present on the legs in both sexes of all three species, with no striking interspecies or intersex differences. Surprisingly, we also found pheromone-related acetate esters in leg extracts of species that lack acetate esters in their female sex pheromone. When we assessed gene expression levels in the leg tissue, we found known and putative pheromone-biosynthesis genes expressed, which suggests that moth legs may be additional sites of pheromone production. To determine possible additional roles of the pheromone compounds on legs, we explored whether these may act as oviposition-deterring signals, which does not seem to be the case. However, when we tested whether these chemicals have antimicrobial properties, we found that two pheromone compounds (16:Ald and 16:OH) reduce bacterial growth. Such an additional function of previously identified pheromone compounds likely coincides with additional selection pressures and, thus, should be considered in scenarios on the evolution of these signals.     

Sex-specific genotype-by-environment interactions for cuticular hydrocarbon expression in decorated crickets, Gryllodes sigillatus: implications for the evolution of signal reliability

Phenotypic traits that convey information about individual identity or quality are important in animal social interactions, and the degree to which such traits are influenced by environmental variation can have profound effects on the reliability of these cues. Using inbred genetic lines of the decorated cricket, Gryllodes sigillatus, we manipulated diet quality to test how the cuticular hydrocarbon (CHC) profiles of males and females respond across two different nutritional rearing environments. There were significant differences between lines in the CHC profiles of females, but the effect of diet was not quite statistically significant. There was no significant genotype-by-environment interaction (GEI), suggesting that environmental effects on phenotypic variation in female CHCs are independent of genotype. There was, however, a significant effect of GEI for males, with changes in both signal quantity and content, suggesting that environmental effects on phenotypic expression of male CHCs are dependent on genotype. The differential response of male and female CHC expression to variation in the nutritional environment suggests that these chemical cues may be under sex-specific selection for signal reliability. Female CHCs show the characteristics of reliable cues of identity: high genetic variability, low condition dependence and a high degree of genetic determination. This supports earlier work showing that female CHCs are used in self-recognition to identify previous mates and facilitate polyandry. In contrast, male CHCs show the characteristics of reliable cues of quality: condition dependence and a relatively higher degree of environmental determination. This suggests that male CHCs are likely to function as cues of underlying quality during mate choice and/or male dominance interactions.     

Acp70A regulates Drosophila pheromones through juvenile hormone induction

Mated Drosophila melanogaster females show a decrease in mating receptivity, enhanced ovogenesis, egg-laying and activation of juvenile hormone (JH) production. Components in the male seminal fluid, especially the sex peptide ACP70A stimulate these responses in females. Here we demonstrate that ACP70A is involved in the down-regulation of female sex pheromones and hydrocarbon (CHC) production. Drosophila G10 females which express Acp70A under the control of the vitellogenin gene yp1, produced fewer pheromones and CHCs. There was a dose-dependent relationship between the number of yp1-Acp70A alleles and the reduction of these compounds. Similarly, a decrease in CHCs and diene pheromones was observed in da > Acp70A flies that ubiquitously overexpress Acp70A. Quantitative-PCR experiments showed that the expression of Acp70A in G10 females was the same as in control males and 5 times lower than in da > Acp70A females.Three to four days after injection with 4.8 pmol ACP70A, females from two different strains, exhibited a significant decrease in CHC and pheromone levels. Similar phenotypes were observed in ACP70A injected flies whose ACP70A receptor expression was knocked-down by RNAi and in flies which overexpress ACP70A N-terminal domain. These results suggest that the action of ACP70A on CHCs could be a consequence of JH activation. Female flies exposed to a JH analog had reduced amounts of pheromones, whereas genetic ablation of the corpora allata or knock-down of the JH receptor Met, resulted in higher amounts of both CHCs and pheromonal dienes.Mating had negligible effects on CHC levels, however pheromone amounts were slightly reduced 3 and 4 days post copulation. The physiological significance of ACP70A on female pheromone synthesis is discussed.     

Myrmecophilous aphids produce cuticular hydrocarbons that resemble those of their tending ants

Aphid-tending ants protect aphids from natural enemies and collect honeydew secreted by the aphids. However, ants also often prey on the aphids they attend. Aphids, therefore, like social parasites of ants, may well have evolved chemical mimicry as an anti-predation strategy. In this study, we aimed to determine whether the aphid Stomaphis yanonis actively produces cuticular hydrocarbons (CHCs) that resemble those of the tending ant Lasius fuji. In the wild, ants put their CHCs on the aphids that they are tending, so in this study we analyzed “ant-free” aphids. Mature aphids that exuviated in the absence of ant attendance had almost all of the hydrocarbon components that the ants’ CHCs had. Moreover, hydrocarbons artificially applied to the aphids’ body surface were lost by exuviation. Taken together, these findings indicate that mature aphids actively produced ant-like CHCs, and they constitute the first documentation of a chemical resemblance between aphids and ants in a specific aphid–ant association.