小蠹虫类异戊二烯类聚集信息素的生物合成

小蠹虫(小蠹科)是重要的森林蛀干害虫,在蛀食坑道时导致树木水分和养分运输系统受到破坏,短时间内对整片森林造成严重的经济危害。聚集信息素在小蠹虫聚集危害过程中扮演着非常重要的角色,目前已有多种小蠹虫聚集信息素成分被鉴定并成功应用于生产防控工作中。类异戊二烯类聚集信息素是小蠹虫中极为重要的一类聚集信息素,其主要成分包括小蠹烯醇、小蠹二烯醇、马鞭草烯醇及其衍生物。本文从类异戊二烯类聚集信息素的生物合成前体物质、生物合成位点、生物合成途径、取食和JHШ调控、微生物与其生物合成关系以及展望6个方面出发,全面阐述了齿小蠹属Ips和大小蠹属Dendroctonus中小蠹虫聚集信息素的生物合成机制及调控机制。文中首先重点阐述了小蠹虫体内以甲羟戊酸途径从头合成小蠹二烯醇以及利用寄主成分α-蒎烯直接合成马鞭草烯醇的生物合成过程;其次阐述了生物合成途径中关键酶和基因对小蠹取食和JHШ处理的响应以及小蠹虫肠道微生物和伴生真菌对该类聚集信息素生物合成的影响;最后,针对小蠹虫类异戊二烯类聚集信息素生物合成研究作了探讨和展望。本文为开发和应用其聚集信息素控制小蠹虫危害提供理论基础。     

蚁形郭公虫生物学研究

蚁形郭公虫Ｔｈａｎａｓｉｍｕｓｆｏｒｍｉｃａｒｉｕｓ（Ｌ．）属鞘翅目（Ｃｈｌａｐｔｅｒａ）,郭公虫科（Ｃｌｅｒｉｄｅａ）,为小蠹虫重要的捕食性天敌［１］。蚁形郭公虫的幼虫和成虫均表现出较强的猎物搜寻和捕食能力,可以借助小蠹虫释放的聚集信息素确定小蠹虫的行踪,并能挖掘隧道深入到树皮内捕食蠹虫［２,３］;此外,该虫的主要捕食时期与多种小蠹虫的发生期相吻合,对春季出现较早的小合虫具有显著的控制效果［４］。９０年代初,为对正在美国大湖地区迅速蔓延的纵坑切梢小蠹实施生物控制,美国农业部特地从欧洲引进了２０…     

云杉八齿小蠹生态学研究进展

云杉八齿小蠹是欧洲最具危害性的小蠹虫之一,嗜食衰弱木,利用雄性产生的信息素进行大面积聚集。聚集信息素不仅具有引诱聚集的作用,其种类和数量还具有调节其聚集密度的效应。在定殖过程中,云杉八齿小蠹既能利用寄主挥发物定位寄主,也能利用非寄主挥发物作为远距离的嗅觉示踪来区别寄主和非寄主。在森林生态系统内捕食性天敌和寄生性天敌很大程度上影响着云杉八齿小蠹的种群密度和行为生态。本文从云杉八齿小蠹的分布与扩散、天敌的种类及控制作用、化学信息物质对小蠹虫及其天敌的行为调控作用以及信息素防治现状等方面进行了综述,并展望了其研究前景。     

小蠧虫防治的方法和措施建议

小蠹虫主要分布在低海拔山区,白天飞出,一般栖息在枯木钻洞,一般很少飞出到巢外。小蠹虫的分布比较广泛、数量也比较大,很容易在林木内密集成群,常年都以树皮或树干为食,很容易造成树木的衰弱,而且会迅速地枯死。本文对小蠹虫的防治方法进行分析和探讨。     

危害松树的小蠹虫与其伴生菌的相互关系

危害健康松属植物的小蠹虫经常与一些特殊的真菌相联系。在小蠹虫危害松属植物的过程中,这些真菌被小蠹虫的一些特殊结构或者体表携带到松属植物上。小蠹虫与其伴生菌的联系表明小蠹虫和其伴生菌之间是一种互惠互利的关系。伴生菌随小蠹虫扩散而被带到新的寄主树木。而伴生菌或作为小蠹虫的食物来源,但更重要的是,有些伴生菌能够通过其菌丝渗透寄主组织,释放毒素,致死寄主树木,以帮助小蠹虫降低寄主抗性。许多研究致力于探索小蠹虫/伴生菌联合体与寄主树木之间关系的特征和确定小蠹虫与其伴生菌相互关系在生态学上的意义。然而,不同小蠹虫和其伴生菌所组成的共生体系,不同小蠹虫的种群数量,和不同环境条件下同种小蠹虫与其伴生菌相互作用方式的差异使我们在研究小蠹虫和其伴生菌这个共生体系时,对它们各自在成功聚集寄主树木过程中所发挥的重要作用的概括变得非常困难。     

危害松树的小蠹虫与其伴生茵的相互关系

危害健康松属植物的小蠹虫经常与一些特殊的真菌相联系.在小蠹虫危害松属植物的过程中,这些真菌被小蠹虫的一些特殊结构或者体表携带到松属植物上.小蠹虫与其伴生菌的联系表明小蠹虫和其伴生菌之间是一种互惠互利的关系.伴生菌随小蠹虫扩散而被带到新的寄主树木.而伴生菌或作为小蠹虫的食物来源,但更重要的是,有些伴生菌能够通过其菌丝渗透寄主组织,释放毒素,致死寄主树木,以帮助小蠹虫降低寄主抗性.许多研究致力于探索小蠹虫/伴生菌联合体与寄主树木之间关系的特征和确定小蠹虫与其伴生菌相互关系在生态学上的意义.然而,不同小蠹虫和其伴生菌所组成的共生体系,不同小蠹虫的种群数量,和不同环境条件下同种小蠹虫与其伴生菌相互作用方式的差异使我们在研究小蠹虫和其伴生菌这个共生体系时,对它们各自在成功聚集寄主树木过程中所发挥的重要作用的概括变得非常困难.     

小蠹虫对针叶类寄主树木的选择危害机制

小蠹虫对寄主植物的选择是一个非常复杂的过程,也是近年来小蠹虫生态学研究的一个热门话题。文章对国内外小蠹虫寄主搜寻、寄主识别,以及适宜寄主选择等机制等方面的最新研究成果进行了系统报道。     

树木-蛀干昆虫-天敌昆虫间的三级营养关系

以小蠹虫和天牛为主的蛀干昆虫是为害林木的重要经济害虫,常对森林生态系统造成重大灾害.本文以林业主要害虫——小蠹虫和天牛为例,综述了树木、蛀干害虫和其天敌组成的三级营养系统间相互关系的研究概况,从不同侧面介绍了近十年来在引诱天敌的信息化学物质的释放部位、活性化合物的鉴定、挥发性化合物的释放动态等方面的研究成果,同时对于天敌对信息化学物质的选择性、天敌视觉在寻找寄主或猎物方面的作用以及是否涉及到影响天敌行为的三级营养以外的其它生物活性物质等的研究进展作了简略介绍.     

小蠹虫人工饲养方法简介

<正> 小蠹虫是重要的森林和检疫害虫之一。其体小,生活隐蔽,繁殖迅速,危害初期很难被人们发现,一旦暴发成灾很难防治,能毁坏大片森林。故我们应加强小蠹虫基本生物学的研究。小蠹虫的人工饲养是研究其生活史、生物学习性、种间和种内的通讯联系、种群的模拟预测模     

大兴安岭地区小蠹虫种类及诱捕器防治研究

本文针对大兴安岭地区的小蠹虫种类进行研究,并通过诱捕器的试验诱集结果,着重研究了小蠹虫的诱捕器防治方法,结果表明落叶松八齿小蠹、云杉八齿小蠹和纵坑切梢小蠹为针叶树的优势种类,诱集结果也证明这3种小蠹的数量占多数,其中落叶松八齿小蠹占绝大多数。     

Individual variation in bark beetle and moth pheromones - a comparison and an evolutionary background

The presently published data of variation in pheromone content in bark beetles (Coleoptera: Scolytidae) and moths (Lepidoptera) are reviewed. In both taxa high coefficients of variation, often up to around 100% for amount of pheromone, seem to be the rule. The contribution to the variation from errors in the chemical methods was small in most studies compared with the intrinsic biological variation. Examples arc given of variation in both absolute amounts, proportions of geometric isomers and in enantiomeric composition. Ratios of isomers often had lower coefficients of variation (25% or less). A lower average variation among sex pheromone in moths could relate to a mating system involving singly calling females, compared with the aggregation pheromone of bark beetles, which often call in large groups. The origin of a resource-based aggregation pheromone system by evolution working via individual selection is outlined.     

Effect of varying monoterpene concentrations on the response of Ips pini (Coleoptera: Scolytidae) to its aggregation pheromone: implications for pest management and ecology of bark beetles

Abstract 1 Host plant terpenes can influence attraction of conifer bark beetles to their aggregation pheromones: both synergistic and inhibitory compounds have been reported. However, we know little about how varying concentrations of individual monoterpenes affect responses. 2 We tested a gradient of ratios of α‐pinene, the predominant monoterpene in host pines in the Great Lakes region of North America, to Ips pini's pheromone, racemic ipsdienol plus lanierone. 3 Ips pini demonstrated a parabolic response, in which low concentrations of α‐pinene had no effect on attraction to its pheromone, intermediate concentrations were synergistic and high concentrations were inhibitory. These results suggest optimal release rates for population monitoring and suppression programmes. 4 Inhibition of bark beetle attraction to pheromones may be an important component of conifer defences. At terpene to pheromone ratios emulating emissions from trees actively responding to a first attack, arrival of flying beetles was low. This may constitute an additional defensive role of terpenes, which are also toxic to bark beetles at high concentrations. 5 Reduced attraction to a low ratio of α‐pinene to pheromone, as occurs when colonization densities become high and the tree's resin is largely depleted, might reflect a mechanism for preventing excessive crowding. 6 Thanasimus dubius, the predominant predator of I. pini, was also attracted to ipsdienol plus lanierone, but its response differed from that of its prey. Attraction increased across all concentrations of α‐pinene. This indicates that separate lures are needed to sample both predators and bark beetles effectively. It also provides an opportunity for maximizing pest removal while reducing adverse effects on beneficial species. This disparity further illustrates the complexity confronting natural enemies that track chemical signals to locate herbivores.     

Fitness consequences of pheromone production and host selection strategies in a tree-killing bark beetle (Coleoptera: Curculionidae: Scolytinae)

Timing of arrival at a resource often determines an individual’s reproductive success. Tree-killing bark beetles can reproduce in healthy trees by attacking in adequate numbers to overcome host defences that could otherwise be lethal. This process is mediated by aggregation and antiaggregation pheromones. Beetles that arrive early in such a “mass attack” must contend with undiminished tree defences, and produce enough pheromones to attract more beetles, but have a head start on gallery construction and egg-laying. Beetles that arrive late may be impeded by competition and diminishing availability of phloem, but should experience fewer costs associated with pheromone production and battling tree defences. We investigated relationships between timing of arrival, body size, pheromone production and fitness in the southern pine beetle, Dendroctonus frontalis. In field experiments, we captured beetles that arrived early (pioneers) and late on slash pine trees, Pinus elliottii, and measured pheromone amounts in their hindguts. We marked gallery entrances of beetles as they landed on a tree and measured their reproductive success after the attack terminated. We found no difference in body size or pheromone amounts between early and late arrivers. Most beetles arrived at the middle of the attack sequence, and excavated longer galleries per day than early arrivers. The number of offspring produced per day by beetles that established galleries midway through mass attack was higher than those that arrived early or very late in the sequence. Our results suggest that beetles do not exhibit adaptive phenotypic plasticity in pre-landing pheromone production, depending on the extent of previous colonisation of a host. Rather, it appears that stabilising selection favours beetles that attack in the middle of the sequence, and contributes to attack synchrony. Synchronous attack on trees is essential before population booms characteristic of tree-killing bark beetles can occur in nature.     

Bark beetles as lidar targets and prospects of photonic surveillance

Forestry is raising concern about the outbreaks of European spruce bark beetle, Ips typographus, causing extensive damage to the spruce forest and timber values. Precise monitoring of these beetles is a necessary step towards preventing outbreaks. Current commercial monitoring methods are catch-based and lack in both temporal and spatial resolution. In this work, light scattering from beetles is characterized, and the feasibility of entomological lidar as a tool for long-term monitoring of bark beetles is explored. Laboratory optical properties, wing thickness, and wingbeat frequency of bark beetles are reported, and these parameters can infer target identity in lidar data. Lidar results from a Swedish forest with controlled bark beetle release event are presented. The capability of lidar to simultaneously monitor both insects and a pheromone plume mixed with chemical smoke governing the dispersal of many insects is demonstrated. In conclusion, entomological lidar is a promising tool for monitoring bark beetles.     

Host resistance elicited by methyl jasmonate reduces emission of aggregation pheromones by the spruce bark beetle, Ips typographus

We treated Norway spruce (Picea abies) stems with methyl jasmonate (MeJA) to determine possible quantitative and qualitative effects of induced tree defenses on pheromone emission by the spruce bark beetle Ips typographus. We measured the amounts of 2-methyl-3-buten-2-ol and (S)-cis-verbenol, the two main components of the beetle's aggregation pheromone, released from beetle entrance holes, along with phloem terpene content and beetle performance in MeJA-treated and untreated Norway spruce logs. As expected, phloem terpene levels were higher and beetle tunnel length was shorter (an indication of poor performance) in MeJA-treated logs relative to untreated logs. Parallel to the higher phloem terpene content and poorer beetle performance, beetles in MeJA-treated logs released significantly less 2-methyl-3-buten-2-ol and (S)-cis-verbenol, and the ratio between the two pheromone components was significantly altered. These results suggest that host resistance elicited by MeJA application reduces pheromone emission by I. typographus and alters the critical ratio between the two main pheromone components needed to elicit aggregation. The results also provide a mechanistic explanation for the reduced performance and attractivity observed in earlier studies when bark beetles colonize trees with elicited host defenses, and extend our understanding of the ecological functions of conifer resistance against bark beetles.     

Pheromone production in bark beetles

The first aggregation pheromone components from bark beetles were identified in 1966 as a mixture of ipsdienol, ipsenol and verbenol. Since then, a number of additional components have been identified as both aggregation and anti-aggregation pheromones, with many of them being monoterpenoids or derived from monoterpenoids. The structural similarity between the major pheromone components of bark beetles and the monoterpenes found in the host trees, along with the association of monoterpenoid production with plant tissue, led to the paradigm that most if not all bark beetle pheromone components were derived from host tree precursors, often with a simple hydroxylation producing the pheromone. In the 1990s there was a paradigm shift as evidence for de novo biosynthesis of pheromone components began to accumulate, and it is now recognized that most bark beetle monoterpenoid aggregation pheromone components are biosynthesized de novo. The bark beetle aggregation pheromones are released from the frass, which is consistent with the isoprenoid aggregation pheromones, including ipsdienol, ipsenol and frontalin, being produced in midgut tissue. It appears that exo-brevocomin is produced de novo in fat body tissue, and that verbenol, verbenone and verbenene are produced from dietary α-pinene in fat body tissue. Combined biochemical, molecular and functional genomics studies in Ips pini yielded the discovery and characterization of the enzymes that convert mevalonate pathway intermediates to pheromone components, including a novel bifunctional geranyl diphosphate synthase/myrcene synthase, a cytochrome P450 that hydroxylates myrcene to ipsdienol, and an oxidoreductase that interconverts ipsdienol and ipsdienone to achieve the appropriate stereochemistry of ipsdienol for pheromonal activity. Furthermore, the regulation of these genes and their corresponding enzymes proved complex and diverse in different species. Mevalonate pathway genes in pheromone producing male I. pini have much higher basal levels than in females, and feeding induces their expression. In I. duplicatus and I. pini, juvenile hormone III (JH III) induces pheromone production in the absence of feeding, whereas in I. paraconfusus and I. confusus, topically applied JH III does not induce pheromone production. In all four species, feeding induces pheromone production. While many of the details of pheromone production, including the site of synthesis, pathways and knowledge of the enzymes involved are known for Ips, less is known about pheromone production in Dendroctonus. Functional genomics studies are under way in D. ponderosae, which should rapidly increase our understanding of pheromone production in this genus. This chapter presents a historical development of what is known about pheromone production in bark beetles, emphasizes the genomic and post-genomic work in I. pini and points out areas where research is needed to obtain a more complete understanding of pheromone production.     

Host tree resistance influencing pheromone production in Ips typographus (Coleoptera: Scolytidae)

Analyses of volatiles in hindguts of Ips typgraphus males from different spruce trees and attack phases are reviewed. The composition of monoterpenes, and the chirality of α-pinene, have been determined in phloem samples. Relationships between compounds emanating from spruce trees and bark beetles, respectively, have been studied. Male beetles depend on their host tree for the production of pinene alcohols. The ratio between the pinene alcohols is almost constant in males boring in the same tree but can vary widely between males from different spruce trees. Very good correlations were found between some host tree monoterpene hydrocarbons and bark beetle produced pinene alcohols. The production of the essential pheromone component 2-methyl-3-buten-2-ol, was not correlated with the monoterpene content in the host trees, while the production of the other essential pheromone component, cis-verbenol, depends on the amount of the precursor, (−)-α-pinene in the phloem. Male beetles boring in a resistant spruce tree will continue to produce the pinene alcohols, including cis -verbenol, as long as the tree defends itself with resin.     

Pine monoterpenes and pine bark beetles: a marriage of convenience for defense and chemical communication

Pine-feeding bark beetles (Coleoptera: Scolytidae) interact chemically with their host pines (Coniferales: Pinaceae) via the behavioral, physiological, and biochemical effects of one class of isoprenoids, the monoterpenes and their derivatives. Pine monoterpenes occur in the oleoresin and function as behaviorally active kairomones for pine bark beetles and their predators, presenting a classic example of tri-trophic chemical communication. The monoterpenes are also essential co-attractants for pine bark beetle aggregation pheromones. Ironically, pine monoterpenes are also toxic physiologically to bark beetles at high vapor concentrations and are considered an important component of the defense of pines. Research over the last 30 years has demonstrated that some bark beetle aggregation pheromones arise through oxygenation of monoterpenes, linking pheromone biosynthesis to the host pines. Over the last 10 years, however, several frequently occurring oxygenated monoterpene pheromone components (e.g., ipsenol, ipsdienol and frontalin) have also been shown to arise through highly regulated de novo pathways in the beetles (reviewed in Seybold and Tittiger, 2003). The most interesting nexus between these insects and their plant hosts involves the late-stage reactions in the monoterpenoid biosynthetic pathway, during which isomeric dimethylallyl diphosphate and isopentenyl diphosphate are ultimately elaborated to stereospecific monoterpenes in the trees and to hydroxylated monoterpenes or bicyclic acetals in the insects. There is signal stereospecificity in both production of and response to the monoterpenoid aggregation pheromones of bark beetles and in response to␣the monoterpenes of the pines. In the California fivespined ips, Ips paraconfusus, we have discovered a number of cytochome P450 genes that have expression patterns indicating that they may be involved in detoxifying monoterpene secondary metabolites and/or biosynthesizing pheromone components. Both processes result in the production of oxygenated monoterpenes, likely with varying degrees of stereospecificity. A behavioral analysis of the stereospecific response of I. paraconfusus to its pheromone is providing new insights into the development of an efficacious bait for the detection of this polyphagous insect in areas outside the western United States. In contrast, a Eurasian species that has arrived in California, the Mediterranean pine engraver, Orthotomicus (Ips) erosus, utilizes both a monoterpenoid (ipsdienol) and a hemiterpenoid (2-methyl-3-buten-2-ol) in its pheromone blend. The stereospecificity of the response of O. erosus to the monoterpenoid appears to be the key factor to the improved potency of the attractant bait for this invasive species.Dedicated to Professor David L. Wood on the occasion of his 75th birthday, January 8, 2006     

Mass attack of trees by Ips typographus induced by sex-specific pheromone: a model of attack dynamics

A model of attack dynamics on a single tree under mass attack by the spruce bark beetle Ips typographus is developed from earlier studies of pheromone production and response of the flying population to the pheromone signal. The model is explored for different numbers and sex-ratios of pioneer beetles and responding beetles. Both numbers of pioneers and responders were important for the dynamics, while the sex-ratio of pioneers was less important. A decreased proportion of males among the responding beetles was important for the result of the model. In the early stage of the attack the model predictions of both numbers and sex-ratio of beetles landing were similar to examples from naturally mass-attacked trees, although the attack proceeded more slowly than in the field. Several aspects of the attack dynamics, such as density-regulation of the number of attacks on the tree, as observed in the field, were not predicted by the model. Gaps in the knowledge of the chemical ecology of the tree-insect relation, such as qualitative change in odours from the tree, spacing mechanisms on the bark surface, and mortality due to host resistence, are pointed out with help of the model.