虫害诱导植物挥发物(HIPVs)对植食性昆虫的行为调控

虫害诱导植物挥发物(herbivore induced plant volatiles,HIPVs)具有植物种类、品种、生育期和部位的特异性,也具有植食性昆虫种类、虫龄、为害程度、为害方式和其他一些环境因子的特异性。由于其释放量明显大于健康植株,因此更易被天敌、害虫以及邻近的植物等所利用,从而调节植物、植食性昆虫与天敌三者之间的相互作用关系,增强植物在自然界的生存竞争能力。本文对HIPVs在植食性昆虫寄主定位行为中的作用、HIPVs对植食性昆虫的种群调控功能及其应用现状2个方面加以综述,并在展望中对目前研究中存在的一些问题进行了探讨。     

虫害诱导挥发物的生态调控功能

虫害诱导挥发物（herbivore-induced plant volatiles, HIPVs）是植物受害虫胁迫后释放的挥发性物质,是植物与周围环境进行信息交流的媒介。环境中的天敌、害虫和植物通过感知HIPVs所携带的信息,对各自的行为或生理生化反应做出相应的调整。介绍了挥发物的种类及主要的生物合成途径,概括了影响天敌依据HIPVs搜寻寄主和猎物的主要因素。综述了这类挥发性物质对植食性昆虫寄主选择或产卵行为的影响,介绍了植物地上部分和地下部分受害后对彼此间接防御的影响,讨论了多种害虫加害同种植物后对天敌搜寻猎物或寄主行为的影响。另外,作为损伤信号,HIPVs还能诱导同株植物未受害部位和邻近植株的防御反应。最后,对HIPVs在害虫防治中的应用现状及前景作了介绍和讨论。     

虫害诱导的植物挥发物:植物与植食性昆虫及其天敌相互作用的进化产物

综述了植物、植食性昆虫及其天敌相互作用的进化过程。虫害诱导的植物挥发物的特征和功能是植物-植食性昆虫-天敌之间长期进化的结果。在植物、植食性昆虫与天敌相互作用的进化过程中,3个不同营养级,包括植物、植食性昆虫和天敌有着各自的调节和利用虫害诱导的植物挥发物的策略。但有一些问题,如通过实验研究得出的诱导防御在田间是否真正能起到保护作用等需进一步研究、阐明。     

虫害诱导的植物挥发物代谢调控机制研究进展

长期受自然界的非生物/生物侵害,植物逐步形成了复杂的防御机制,为防御植食性昆虫的为害,植物释放虫害诱导产生的挥发性化合物(herbivore-induced plant volatiles,HIPVs)。HIPVs是植物-植食性昆虫-天敌三级营养关系之间协同进化的结果。HIPVs的化学组分因植物、植食性昆虫种类的不同而有差异。生态系统中,HIPVs可在植物与节肢动物、植物与微生物、虫害植物与邻近的健康植物、或同一植株的受害和未受害部位间起作用,介导防御性反应。HIPVs作为寄主定位信号,在吸引捕食性、寄生性天敌过程中起着重要作用。HIPVs还可以作为植物间信息交流的工具,启动植株的防御反应而增强抗虫性。不论从生态学还是经济学角度来看,HIPVs对于农林生态系中害虫综合治理策略的完善具有重要意义。前期的研究在虫害诱导植物防御的化学生态学方面奠定了良好基础,目前更多的研究转向阐述虫害诱导植物抗性的分子机制。为了深入了解HIPVs的代谢调控机制,主要从以下几个方面进行了综述。因为植食性昆虫取食造成的植物损伤是与昆虫口腔分泌物共同作用的结果,所以首先阐述口腔分泌物在防御反应中的作用。挥发物诱导素volicitin和β-葡萄糖苷酶作为口腔分泌物的组分,是产生HIPVs的激发子,通过调节伤信号诱发HIPVs的释放。接着阐述了信号转导途径对HIPVs释放的调节作用,并讨论了不同信号途径之间的交互作用。就HIPVs的代谢过程而言,其过程受信号转导途径(包括茉莉酸、水杨酸、乙烯、过氧化氢信号途径)的调控,其中茉莉酸信号途径是诱发HIPVs释放的重要途径。基于前人的研究,综述了HIPVs的主要代谢过程及其过程中关键酶类的调控作用。文中的HIPVs主要包括萜烯类化合物、绿叶挥发物和莽草酸途径产生的芳香族化合物,如水杨酸甲酯和吲哚等。作为化学信号分子,这些化合物中的一部分还能激活邻近植物防御基因的表达。萜烯合酶是各种萜烯类化合物合成的关键酶类,脂氧合酶、过氧化氢裂解酶也是绿叶挥发物代谢途径中的研究热点,而苯丙氨酸裂解酶和水杨酸羧基甲基转移酶分别是合成水杨酸及其衍生物水杨酸甲酯的关键酶类。这些酶类的基因在转录水平上调控着HIPVs代谢途径。最后展望了HIPVs的研究前景。     

虫害诱导的植物挥发物:基本特性、生态学功能及释放机制

植物在遭受植食性昆虫攻击时,能通过释放挥发物调节植物、植食性昆虫及其天敌三者间的相互关系,并由此而防御植食性昆虫。主要就虫害诱导的植物挥发物的基本特性、生态学功能及其释放机制进行了系统性综述,并提出了今后的研究方向。     

虫害诱导植物挥发物（HIPVs）：从诱导到生态功能

植物被植食性昆虫危害后,其地上和地下部分均能释放虫害诱导植物挥发物(HIPVs).HIPVs不仅能招引植食性昆虫的天敌,而且还可对植食性昆虫和临近植物产生影响,从而调节植物、植食性昆虫、天敌三者之间的相互关系.根据最近的研究结果,主要就HIPVs的诱导物类型、虫害后植物体内的早期信号和信号传导、HIPVs的合成、释放、组成,以及其生态功能等进行了系统性的综述,并提出了今后该领域的研究方向.     

植食性昆虫诱导的挥发物及其在植物通讯中的作用

正常健康植株的挥发物代谢维持在基底水平,当遭到昆虫取食时,植物释放出特定的挥发物,用来招引害虫的天敌,还能诱导邻近植株产生防御反应.文章就此问题的研究进展作了介绍.     

植物创伤诱导的挥发物及其信号功能

植物在受到动物或病原侵害时,会释放特异的挥发作作为信号,这些信号分子是植物防御系统的重要组成部分,文章对植物创伤诱导挥发物的种类,诱导因子及生理作用等的研究进展,以及这一领域的研究潜力和发展方向作了评述。     

虫害诱导的水稻挥发物对褐飞虱的驱避作用

水稻叶片在遭受害虫的危害时,释放一些特异性的挥发性物质。这些挥发性信息物质在调节植物、植食性昆虫及其天敌的相互关系中有重要作用。利用“Y”型嗅觉仪研究了褐飞虱对虫害诱导的11种水稻挥发物（浓度为2 Μl 化合物溶解在100 Μl丙酮溶液中）的行为反应。结果表明,(E)-2-己烯醛、(E)-2-己烯-1-醇、-2-庚醇和水杨酸甲酯等4种化合物对褐飞虱成虫有显著的驱避作用,而(Z)-3-己烯-1-醇、-2-庚酮、柠檬烯、罗勒烯、芳樟醇、-β-子丁香烯和橙花叔醇等7种物质对褐飞虱成虫的选择行为无显著影响。浓度梯度的实验表明,浓度(不同体积的待测化合物用100 Μl丙酮稀释)较低的(E)-2-己烯醛（0.5 Μl、1 Μl）和芳樟醇（05 Μl、1 Μl、5 Μl）对褐飞虱成虫的选择行为无显著影响,而较高浓度的(E)-2-己烯醛（5 Μl、10 Μl）和芳樟醇（10 Μl）对褐飞虱成虫均有明显的驱避作用。     

新旧创伤诱导的不同挥发物对寄生蜂选择性的影响

植物在植食性昆虫危害的压力下会释放出化合物,而寄生蜂却可以利用这些诱导出的植物挥发物定位寄主。植物叶表面受到创伤都会释放出最常见的绿叶挥发物（green leaf volatiles）,而植食性昆虫取食引诱的挥发物会因害虫和受害植物的不同而异。植物在受到害虫造成的创伤后会马上挥发出绿叶性化合物,而一些萜稀类化合物（terpenoids）只有在旧的创伤部位才会大量挥放。那么不同的寄生蜂可以识别这些化合物吗？     

Combined use of herbivore‐induced plant volatiles and sex pheromones for mate location in braconid parasitoids

Herbivore‐induced plant volatiles (HIPVs) are important cues for female parasitic wasps to find hosts. Here, we investigated the possibility that HIPVs may also serve parasitoids as cues to locate mates. To test this, the odour preferences of four braconid wasps – the gregarious parasitoid Cotesia glomerata (L.) and the solitary parasitoids Cotesia marginiventris (Cresson), Microplitis rufiventris Kokujev and Microplitis mediator (Haliday) – were studied in olfactometers. Each species showed attraction to pheromones but in somewhat different ways. Males of the two Cotesia species were attracted to virgin females, whereas females of M. rufiventris were attracted to virgin males. Male and female M. mediator exhibited attraction to both sexes. Importantly, female and male wasps of all four species were strongly attracted by HIPVs, independent of mating status. In most cases, male wasps were also attracted to intact plants. The wasps preferred the combination of HIPVs and pheromones over plant odours alone, except M. mediator, which appears to mainly use HIPVs for mate location. We discuss the ecological contexts in which the combined use of pheromones and HIPVs by parasitoids can be expected. To our knowledge, this is the first study to show that braconid parasitoids use HIPVs and pheromones in combination to locate mates.     

Biological control of Myzus persicae (Hemiptera: Aphididae) through banker plant system in protected crops

Banker plants with Aphidius colemani were tested in greenhouse for control of Myzus persicae on arugula and sweet pepper crops and compared to inoculative releases of parasitoids. Banker plants system consisted of pots of oat (non-crop plant) infested with Rhopalosiphum padi (non-pest herbivore). The non-pest herbivore serves as an alternative host for A. colemani (parasitoid of the target crop pest). In the arugula crop significant differences in the pest population between the two strategies of biological control showed the lowest densities of the pest when introducing the banker plant system. In the sweet pepper crop, there was no difference in the pest population between the two strategies of biological control.     

硅对植物抗虫性的影响及其机制

硅不是植物必需营养元素,但硅在提高植物对一系列非生物和生物胁迫的抗性方面都具有重要作用。综述了硅对植物抗虫性的影响及其机制。在多数植物中,增施硅肥可增强其抗虫性;所增强的抗性与硅肥种类和施用方式之间存在关系。植物组织中沉积的硅可增加其硬度和耐磨度,降低植物可消化性,从而增强植物组成性防御,包括延缓昆虫生长发育、降低繁殖力、减轻植物受害程度;植物体内的硅含量以及硅沉积的位点和排列方式影响组成性防御作用的强度。此外,硅可以调节植物诱导性防御,包括直接防御和间接防御,直接防御涉及增加有毒物质含量、产生局部过敏反应或系统获得抗性、产生有毒化合物和防御蛋白,从而延缓昆虫发育;间接防御主要通过释放挥发性化合物吸引植食性昆虫的捕食性和寄生性天敌而导致植食性昆虫种群下降。     

Advances in research of induced resistance to insects in cotton

Any change in a plant that occurs following herbivory or environmental factors is an induced response. These changes include phytochemical induction, increases in physical defenses, emission of volatiles that attract predators and parasitoids of herbivores, and reduction in plant nutritional quality for herbivores, which is termed induced resistance. Induced resistance has been demon-strated ubiquitously in plants. It is one of our goals to review what is known about the induced resistance to herbivorous insects in cotton, including three resistance secondary metabolites (terpenoid, tannin, and flavonoids) that are contained at any significant levels of resistance to herbivorous insects in cotton cultivates. In many cases, the quantities or quality of secondary metabolites in plant are changed after attacked by insects. This review focuses on induced plant resistance as quantitative or qualitative enhancement of defense mechanism against insect pests, especially on the abiotic-elicitors-induced resistance in cotton plants. The abiotic-elicitor of cupric chloride, an exogenous inorganic compound, may induce the second-ary metabolites accumulation and is referred to as a copper-inducible elicitor (CIE). Finally, we discuss how copper-inducible elicitor may be used in the Integrated Pest Management (IPM) system for cotton resistance control.     

Phylogenetic test of speciation by host shift in leaf cone moths (Caloptilia) feeding on maples (Acer)

The traditional explanation for the exceptional diversity of herbivorous insects emphasizes host shift as the major driver of speciation. However, phylogenetic studies have often demonstrated widespread host plant conservatism by insect herbivores, calling into question the prevalence of speciation by host shift to distantly related plants. A limitation of previous phylogenetic studies is that host plants were defined at the family or genus level; thus, it was unclear whether host shifts predominate at a finer taxonomic scale. The lack of a statistical approach to test the hypothesis of host‐shift‐driven speciation also hindered studies at the species level. Here, we analyze the radiation of leaf cone moths (Caloptilia) associated with maples (Acer) using a newly developed, phylogeny‐based method that tests the role of host shift in speciation. This method has the advantage of not requiring complete taxon sampling from an entire radiation. Based on 254 host plant records for 14 Caloptilia species collected at 73 sites in Japan, we show that major dietary changes are more concentrated toward the root of the phylogeny, with host shift playing a minor role in recent speciation. We suggest that there may be other roles for host shift in promoting herbivorous insect diversification rather than facilitating speciation per se.     

Advances in research of induced resistance to insects in cotton

Any change in a plant that occurs following herbivory or environmental factors is an induced response. These changes include phytochemical induction, increases in physical defenses, emission of volatiles that attract predators and parasitoids of herbivores, and reduction in plant nutritional quality for herbivores, which is termed induced resistance. Induced resistance has been demonstrated ubiquitously in plants. It is one of our goals to review what is known about the induced resistance to herbivorous insects in cotton, including three resistance secondary metabolites (terpenoid, tannin, and flavonoids) that are contained at any significant levels of resistance to herbivorous insects in cotton cultivates. In many cases, the quantities or quality of secondary metabolites in plant are changed after attacked by insects. This review focuses on induced plant resistance as quantitative or qualitative enhancement of defense mechanism against insect pests, especially on the abiotic-elicitors-induced resistance in cotton plants. The abiotic-elicitor of cupric chloride, an exogenous inorganic compound, may induce the secondary metabolites accumulation and is referred to as a copperinducible elicitor (CIE). Finally, we discuss how copperinducible elicitor may be used in the Integrated Pest Management (IPM) system for cotton resistance control.     

Roles of plant volatiles in defence against microbial pathogens and microbial exploitation of volatiles

Plants emit a large variety of volatile organic compounds during infection by pathogenic microbes, including terpenes, aromatics, nitrogen‐containing compounds, and fatty acid derivatives, as well as the volatile plant hormones, methyl jasmonate, and methyl salicylate. Given the general antimicrobial activity of plant volatiles and the timing of emission following infection, these compounds have often been assumed to function in defence against pathogens without much solid evidence. In this review, we critically evaluate current knowledge on the toxicity of volatiles to fungi, bacteria, and viruses and their role in plant resistance as well as how they act to induce systemic resistance in uninfected parts of the plant and in neighbouring plants. We also discuss how microbes can detoxify plant volatiles and exploit them as nutrients, attractants for insect vectors, and inducers of volatile emissions, which stimulate immune responses that make plants more susceptible to infection. Although much more is known about plant volatile–herbivore interactions, knowledge of volatile–microbe interactions is growing and it may eventually be possible to harness plant volatiles to reduce disease in agriculture and forestry. Future research in this field can be facilitated by making use of the analytical and molecular tools generated by the prolific research on plant–herbivore interactions.     

Caterpillar‐induced plant volatiles attract conspecific herbivores and a generalist predator

Plants release volatiles in response to caterpillar feeding that attracts natural enemies of the herbivores, a tritrophic interaction which has been considered to be an indirect plant defence against herbivores. On the other hand, the caterpillar‐induced plant volatiles have been reported to either repel or attract conspecific adult herbivores. This work was undertaken to investigate the response of both herbivores and natural enemies to caterpillar‐induced plant volatiles in apple orchards. We sampled volatile compounds emitted from uninfested apple trees, and apple trees infested with generalist herbivore the pandemis leafroller moth, Pandemis pyrusana (Lepidoptera, Tortricidae) larvae using headspace collection and analysed by gas chromatography/mass spectrometry. Infested apple trees uniquely release six compounds (benzyl alcohol, phenylacetonitrile, phenylacetaldehyde, 2‐phenylethanol, indole and (E)‐nerolidol). These compounds were tested on two species of herbivores and one predator in apple orchards. Binary blends of phenylacetonitrile + acetic acid or 2‐phenylethanol + acetic acid attracted a large number of conspecific male and female adult herbivores. The response of pandemis leafroller to herbivore‐induced plant volatiles (HIPVs) was so pronounced that over one thousand and seven hundred conspecific male and female adult herbivores were caught in traps baited with HIPVs in three‐day trapping period. In addition, significantly higher number of male and female obliquebanded leafroller, Choristoneura rosaceana (Lepidoptera, Tortricidae), was caught in traps baited a binary blend of 2‐phenylethanol + acetic acid, or a ternary blend contains 2‐phenylethanol and phenylacetonitrile + acetic acid. This result challenges the current paradigm hypothesized that HIPVs repel herbivores and question the indirect defensive function proposed for these compounds. On the other hand, a ternary blend of phenylacetonitrile and 2‐phenylethanol + acetic acid attracted the largest numbers of the general predator, the common green lacewing, Chrysoperla plorabunda. To our knowledge, this is the first record of the direct attraction of conspecific adult herbivores as well as a predator to the caterpillar‐induced plant volatiles in the field.     

Plant-parasitic nematodes affecting vegetable crops in greenhouses

This work focuses on investigating plant-parasitic nematodes that affect greenhouse vegetables. The study took place in the Rostov region (Russian Federation) between May 2019 and May 2020 and involved 180 samples of soil and roots of 30 different vegetables in the families Cucurbitaceae (6), Solanaceae (8), Umbelliferae (8), Lamiaceae (4) and Allioideae (4) from 20 intensive farming locations. In this study, 11 nematode genera were detected. The most common genus was Meloidogyne, followed by Helicotylenchus, Pratylenchus, and Scutellonema. The highest Meloidogyne densities were detected in cucumbers, green peppers, carrots, eggplants, basil, and celery. Onions were not infected with Meloidogyne at all. Plant diseases caused by Pratylenchus, Scutellonema and Helicotylenchus were present in 29.7%, 51.5% and 81.6% of all crops examined, respectively. Xiphinema were found exclusively in carrots and celery, while Ditylenchus were only present in tomatoes and carrots (for each, the prevalence was 2.1%). The relative abundance of Meloidogyne, Helicotylenchus, and Pratylenchus was 58.3%, 10.4%, and 2.1%, respectively. As regards other genera, the relative abundance was less than 1%. The results show that soil properties are as important for the abundance, distribution and structure of the plant-parasitic nematode communities as the host plant. Findings may be helpful in improving the vegetable pest controls.     

Plant volatiles moderate response to aggregation pheromone in Colorado potato beetle

Abstract:  The orientation of the Colorado potato beetle, Leptinotarsa decemlineata , to a male-produced aggregation pheromone, ( S )-3,7-dimethyl-2-oxo-oct-6-ene-1,3-diol, a three-component plant attractant blend [comprised of ( Z )-3-hexenyl acetate + (±)-linalool + methyl salicylate], and other potato volatiles (nonanal and 2-phenylethanol) were tested. All compounds were previously shown to be active in coupled gas chromatography/electroantennogram experiments. Both the three-component plant attractant blend and 2-phenylethanol were attractive to adult beetles. While male beetles oriented preferentially to both plant attractants vs. a control, females showed little preference. Combining the plant attractants with the pheromone resulted in sexually dimorphic responses similar to those seen with either plant attractant alone. Addition of nonanal abolished the sexually dimorphic response to the pheromone + 2-phenylethanol blend; the new three-component blend was attractive to both sexes. In both laboratory bioassays and field experiments, a combination of the pheromone + the three-component plant attractant was preferred over the plant attractant alone. Thus, it seems likely that combinations of pheromone + plant volatiles may be the most efficacious for field use.