大蜂螨对一些气味物质的趋性

狄斯瓦螨Varroa destructor Anderson & Trueman是意大利蜜蜂Apis mellifera Spinola的主要外寄生螨。雌成螨在幼虫巢房封盖前不久侵入幼虫巢房,并开始繁殖为害。从雌成螨在一个很短的时间内进入蜜蜂幼虫巢房,以及雄蜂幼虫巢房蜂螨的寄生率明显高于工蜂幼虫巢房的现象,表明蜜蜂幼虫体表一些信息素(semiochemicals)可能起着重要的引诱作用。作者对与大蜂螨相关的19种气味物质进行筛选,并对封盖前工蜂幼虫和雄蜂幼虫表皮挥发物进行气谱及气-质联谱测定。结果表明:雄蜂6龄幼虫对大蜂螨的引诱作用显著高于丁香水等10种气味物质。工蜂和雄蜂末龄幼虫体表挥发物的共有组份是9-二十三烯(C23H46),但它在雄蜂幼虫中所占的比例要明显高于工蜂幼虫。工蜂幼虫的特有主要组分是十八烷(C18H38)和9-甲基十九烷(C19H40);而雄蜂幼虫的特有主要组分是二十五烷(C25H52)和二十三烷(C23H48)。     

狄斯瓦螨和蜜蜂病毒对意大利蜜蜂蜂群越冬的影响

越冬期是蜂群损失最主要的阶段.通过比较分析45个意大利蜜蜂Apis mellifera ligustica蜂群在繁殖越冬蜂前的狄斯瓦螨Varroa destructor寄生率和病毒感染情况、越冬表现及越冬期存活蜂群的病毒感染情况等,探究与越冬期蜜蜂健康紧密相关的影响因素.结果表明,繁殖越冬蜂前蜂群的狄斯瓦螨寄生率与蜜蜂...     

Potential mechanism for detection by Apis mellifera of the parasitic mite Varroa destructor inside sealed brood cells

Abstract The parasitic mite Varroa destructor Anderson & Trueman is a major pest of the honeybee Apis mellifera L. throughout the world. Chemical agents currently used for mite control leave contaminating residues and promote pesticide resistance. As an alternative means of control, it would be useful to identify natural substances enabling bees to detect Varroa inside brood cells. These substances could then be used to trigger mite hygienic behaviour by bees.
In this study several techniques were used to screen substances that might allow detection of infested brood cells by bees. Gas chromatography-mass spectrometry analysis was performed on substances extracted in dichloromethane from the contents of brood cells. Solid phase microextraction and solid injection were performed on substances obtained from living and dead Varroa, respectively. Electroantennography was performed to assess the sensitivity of olfactory receptors in bee antennae to some of these substances.
Principal component analysis based on proportions of cuticular substances allowed discrimination between bees and other cell contents. Foundress Varroa exhibited the greatest dissimilarity to healthy pupae that were used as controls. Immature Varroa and faecal material were intermediate. High molecular weight compounds, mainly dimethylalkanes, were proportionally the most characteristic components of foundress Varroa . This finding suggests that these compounds would be the most apt to induce uncapping of cells infested by Varroa . Solid-phase microextraction and solid injection demonstrated the presence of aliphatic acids, esters, and one alcohol, eicosenol, in Varroa . Electroantennographic recordings showed that mite-resistant bees were more responsive to some acids and one ester. We speculate that these compounds may be involved in recognition of living Varroa by honeybees.     

蜜蜂寄生螨—狄斯瓦螨的研究进展

狄斯瓦螨Varroa destructor是对世界养蜂业危害最大的蜜蜂寄生虫,严重危害蜜蜂封盖幼虫、蛹和成蜂,并携带和传播蜜蜂病毒,造成蜂群生产力严重下降乃至全群毁灭。狄斯瓦螨的有效防治措施的研发有赖于对其研究进展的了解,本文综述了以下3方面的研究概况:1)狄斯瓦螨的繁殖特性;2)对蜜蜂的危害;3)主要防治方法。以期为蜂螨相关的研究和应用奠定基础。     

微孢子虫和狄斯瓦螨分别侵染后的意蜂血淋巴蛋白质含量变化

微孢子虫Nosema apis和狄斯瓦螨微孢子虫Nosema apis和狄斯瓦螨 Varroa destructor (Acari: Varroidae)均为危害意蜂Apis mellifera的重要寄生虫,该文对其危害后意蜂血淋巴蛋白质含量的变化进行了研究。用考马斯亮蓝法测定了意蜂受侵染后血淋巴的蛋白质总量,并用高压超薄层等电点聚焦法进行血淋巴蛋白质分类。结果显示,病蜂血淋巴蛋白质总量,在人工感染微孢子虫后1～10天呈微孢子虫Nosema apis和狄斯瓦螨 Varroa destructor (Acari: Varroidae)均为危害意蜂Apis mellifera的重要寄生虫,该文对其危害后意蜂血淋巴蛋白质含量的变化进行了研究。用考马斯亮蓝法测定了意蜂受侵染后血淋巴的蛋白质总量,并用高压超薄层等电点聚焦法进行血淋巴蛋白质分类。结果显示,病蜂血淋巴蛋白质总量,在人工感染微孢子虫后1～10天呈上升趋势,然后逐渐下降,感染后12～27天保持在感染前意蜂血淋巴总蛋白质含量水平以下。螨侵染后意蜂血淋巴蛋白质含量明显增高,与健康意蜂相比差异极显著。高压超薄层等电点聚焦分析表明：狄斯瓦螨自然侵染意蜂后,意蜂血淋巴蛋白质组分与健康对照组相比发生了明显改变。这些结果提示,意蜂对于微孢子虫或狄斯瓦螨的侵染产生了一定的免疫反应。     

Varroa management in small bites

Chelifers (Arachnida: Pseudoscorpionida), also known as pseudoscorpions, have been reported to be beneficial honeybee hive generalist pest predators for over 100 years and are occasionally noted by beekeepers in their hives. We collected chelifers within or closely associated with beehives in New Zealand. Under video observation conditions, they predated upon varroa mites while studiously ignoring bee larvae. Varroa mites reproduce at exponential rates during the spring season, and current chemical miticides rely on single treatments aiming for at least 90% control. An alternate strategy, removal of mites at a rate matching their reproductive capacity, although mathematically obvious, fails unless a suitable biological control agent is available. Our observations build on over 100 years of sporadic work to provide further evidence that chelifers show clear potential to be a suitable predator for varroa management in beehives. Approximately 25 chelifers can be expected to manage varroa populations in a single hive.     

狄斯瓦螨和蜜蜂残翅病毒对蜜蜂健康的协同影响

世界各地大范围的西方蜜蜂Apis mellifera蜂群损失现象已引起科学界和公众的持续关注。狄斯瓦螨Varroa destructor和蜜蜂残翅病毒(Deformed wing virus,DWV)是西方蜜蜂群中最主要的两大生物威胁。尽管二者侵害蜜蜂均已有较长历史,但直至近十年来的研究才发现两者间的协同效应对蜂群健康的影响远超过其单独作用时所造成的危害:(1)蜜蜂残翅病毒可在狄斯瓦螨体内大量复制,继而进一步传播;(2)狄斯瓦螨的刺吸行为使病毒粒子跨越寄主的生理屏障而直接进入蜜蜂血淋巴;(3)狄斯瓦螨的寄生促使蜜蜂残翅病毒的高毒力毒株在蜂群中优势扩增和盛行;(4)狄斯瓦螨影响蜜蜂个体发育与免疫系统等生理机能,以致降低了蜂群对病毒的抵抗力;(5)蜜蜂残翅病毒对宿主造成的免疫抑制有利于狄斯瓦螨的寄生与繁殖。狄斯瓦螨、蜜蜂残翅病毒和西方蜜蜂间的关系已经成为昆虫外寄生物、病原体与寄主相互作用研究的一个典型模型。本文对近十年该领域的相关研究进行综述,以期为蜂群损失的原因调查以及昆虫寄生虫、病原微生物与寄主间关系的研究提供参考和借鉴。     

Fatty acid composition of the parasitic mite Varroa destructor and its host the worker prepupae of Apis mellifera

The present study analyzes the fatty acid (FA) profile of lipids isolated from Varroa destructor Anderson & Trueman, a parasitic mite of the honey bee (Apis mellifera L.), uninfected and infected worker prepupae of the Carnolian subspecies Apis mellifera carnica Pollmann, and bee bread fed to the worker brood. Significant differences are observed in the FA profiles of lipids isolated from parasites, hosts and bee bread. Parasitism by V. destructor (henceforth, varroosis) induces visible changes in the lipid profile of worker prepupae. In infected prepupae, the percentage of total saturated FAs is lower and the percentage of unsaturated FAs is higher than in uninfected insects. These differences result from significant changes in the percentages of FAs that are most abundant in the evaluated groups (i.e. C16:0, C18:1 9c, C18:2n‐6 and C18:3n‐3 FAs). In mites and in uninfected and infected prepupae, the predominant FAs are oleic acid (41.07 ± 2.26%, 42.79 ± 1.21% and 45 ± 0.20%, respectively) and palmitic acid (22.62 ± 0.87%, 39.48 ± 0.43% and 36.84 ± 0.22%, respectively). Highly significant differences in FA composition are noted between bee bread and worker brood. The results suggest specific mechanisms of FA uptake, accumulation and metabolism in the food chain of this parasitic association, beginning from the food processed by nurse bees for larval feeding, through host organisms (worker brood) to V. destructor mites.     

Association of Varroa destructor females in multiply infested cells of the honeybee Apis mellifera

The genetic diversity of Varroa destructor (Anderson &Trueman)is limited outside its natural range due to population bottlenecks and its propensity to inbreed.In light of the arms race between V.destructor and its honeybee (Apis mellifera L.)host, any mechanism enhancing population admixture of the mite may be favored.One way that admixture can occur is when two genetically dissimilar mites coinvade a brood cell, with the progeny of the foundresses admixing.We determined the relatedness of 393 pairs of V.destructor foundresses,each pair collected from a single bee brood cell (n =five colonies).We used six microsatellites to identify the genotypes of mites coinvading a cell and calculated the frequency of pairs with different or the same genotypes.We found no deviation from random coinvasion,but the frequency of cells infested by mites with different genotypes was high.This rate of recombination,coupled with a high transmission rate of mites,homogenized the allelic pool of mites within the apiary.     

Resistance of the honey bee, Apis mellifera to the acarian parasite Varroa destructor: behavioural and electroantennographic data

Abstract. One way in which Apis mellifera honey bees resist Varroa destructor is by detection and elimination of nestmates. This study uses behavioural tests and electroanntennography to assess the role of chemostimuli in recognition by honey bees of this acarian ectoparasite. Behavioural tests using living or dead parasites involved observation of honey bee grooming activity (antennation) under controlled conditions in Petri dishes, and removal behaviour (uncapping and elimination of parasitized and unparasitized control brood cells) under natural conditions. Some bees from colonies with both small and large parasite populations showed aggressive behaviour (biting). No difference was observed according to whether the mite was dead or alive. Under natural conditions, bees uncapped more parasitized cells than control cells. Electroantennographic tests were performed to measure sensitivity to various Varroa extracts at three concentrations (10, 20 and 30 Varroa Equivalents). Only 30 Varroa Equivalent methanol extracts made from Varroa collected from brood cells elicited significantly greater antennal response than controls (pure solvent). All three methanol extracts elicited significantly greater antennal response than controls. No response was observed using Varroa extracts made with acetone or hexane. These findings suggest that polar products may act as chemostimuli for recognition of V. destructor by honey bees. Further study will be necessary to determine which polar products are involved in this recognition and assess grooming and removal behaviour using these products.     

蜜蜂巢房大小影响狄斯瓦螨的繁殖行为

在具有相同类型幼虫的雄蜂和工蜂巢房中,人工接入狄斯瓦螨Varroa destructorAnderson&Trueman,比较巢房大小不同,对于螨繁殖的影响。结果显示:狄斯瓦螨在具有工蜂幼虫的工蜂房(WW)中的繁殖率为94.4%,而在具有工蜂幼虫的雄蜂房(WD)中繁殖率只有27.7%,差异极显著。在具有工蜂幼虫的工蜂房中,每只雌螨产出后代的平均数为3.35±1.56只;在具有工蜂幼虫的雄蜂房中每只雌螨产出后代的平均数为0.49±0.93只,差异极显著。表明:在具有相同类型幼虫存在的情况下,狄斯瓦螨喜欢较小的巢房,狄斯瓦螨在较小巢房中的繁殖能力明显高于较大的巢房。     

Patterns in Varroa destructor depend on bee host abundance,availability of natural resources,and climate in Mediterranean apiaries

1. Varroa destructor Linnaeus (Acari: Varroidae) is one of the greatest threats to apiculture. This study examines the role of host density, natural resource availability for bees, the management and climate in driving spatial and annual variability in the abundance of Varroa, and the occurrence of colony losses, recorded in apiaries across a Mediterranean island over a 2‐year period, using a hierarchical generalised linear model framework. 2. The seasonal abundance of Varroa showed a bimodal pattern with two peaks, the first one being in spring and larger than the second one located in summer/autumn. In contrast, bee colony losses were mainly concentrated in autumn/winter. 3. The abundance patterns of Varroa were best explained by models combining host, climate, and resource availability. A key novel finding was that low availability of natural flowering resources leads to high levels of infestation of Varroa, highlighting the importance of preserving natural resources around apiaries for the maintenance of pollination services in the landscape. Varroa abundance was also found to increase as bee density increased, probably as a result of the greater brood availability. Moreover, Varroa abundance increased as temperatures decrease and decreases as relative humidity increases, which is consistent with previous studies. Anti‐varroa treatments were only found to impact Varroa levels in the second 6 months of the year. Organic treatments outperformed synthetic treatments. 4. Empirical research on optimal seasonal timing and combinations of treatments, as well as impacts of climate and resource availability on natural dynamics of bees and Varroa in different climate zones, is urgently required.     

钠离子通道与蜜蜂狄斯瓦螨对氟胺氰菊酯的抗性机理

狄斯瓦螨Varroadestructor是全世界蜜蜂最严重的寄生虫 ,目前 ,它对主要防治药物———拟除虫菊酯类的氟胺氰菊酯已产生明显抗性 ,严重影响其防治效果。近年来神经生理学研究结果证实 :电压门控的钠离子通道是拟除虫菊酯作用的位点。钠通道结构的改变 ,是拟除虫菊酯类杀虫剂毒理的主要基础 ,也是产生抗药性的基础。该文介绍了近年来国内外研究电压门控钠离子通道、拟除虫菊酯对钠通道的作用、钠通道与拟除虫菊酯的抗性和狄斯瓦螨对氟胺氰菊酯抗性机理研究的新进展     

Tyramine functions as a toxin in honey bee larvae during Varroa-transmitted infection by Melissococcus pluton

From wounds of honey bee pupae, caused by the mite Varroa destructor, coccoid bacteria were isolated and identified as Melissococcus pluton. The bacterial isolate was grown anaerobically in sorbitol medium to produce a toxic compound that was purified on XAD columns, gelfiltration and preparative HPLC. The toxic agent was identified by GC-MS and FTICR-MS as tyramine. The toxicity of the isolated tyramine was tested by a novel mobility test using the protozoon Stylonychia lemnae. A concentration of 0.2 mg/ml led to immediate inhibition of mobility. In addition the toxicity was studied on honey bee larvae by feeding tyramine/water mixtures added to the larval jelly. The lethal dosis of tyramine on 4-5 days old bee larvae was determined as 0.3 mg/larvae when added as a volume of 20 microl to the larval food in brood cells. Several other biogenic amines, such as phenylethylamine, histamine, spermine, cadaverine, putrescine and trimethylamine, were tested as their hydrochloric salts for comparison and were found to be inhibitory in the Stylonychia mobility test at similar concentrations. A quantitative hemolysis test with human red blood cells revealed that tyramine and histamine showed the highest membranolytic activity, followed by the phenylethylamine, trimethylamine and spermine, while the linear diamines, cadaverine and putrescine, showed a significantly lower hemolysis when calculated on a molar amine basis. The results indicate that tyramine which is a characteristic amine produced by M. pluton in culture, is the causative agent of the observed toxic symptoms in bee larvae. Thus this disease, known as European foulbrood, is possibly an infection transmitted by the Varroa destructor mite.     

信息素介导的西方蜜蜂与狄斯瓦螨互作研究进展

狄斯瓦螨是一种严重危害西方蜜蜂的体外寄生螨,是世界养蜂业的最大威胁.人们广泛采用化学方法防治狄斯瓦螨,但易引起狄斯瓦螨的抗药性、蜜蜂中毒和蜂产品药物残留等问题.为此,人们尝试了多种蜂螨绿色防控技术.其中利用蜜蜂信息素防治狄斯瓦螨是一个重要的研究方向.研究表明,狄斯瓦螨能利用蜜蜂信息素识别处于不同发育阶段的寄主,并对特定时期的寄主表现出高度的选择性.近年来,多种能作用于狄斯瓦螨的蜜蜂信息素相继被报道.这些信息素包括成蜂、蛹和幼虫信息素.有的信息素对狄斯瓦螨表现出驱避作用,有的则表现出引诱作用.本文对这些信息素的种类、主要组成成分、对狄斯瓦螨的作用等进行了综述,旨在为今后的研究与应用提供参考.     

Behavioural responses of Varroa destructor (Acari: Varroidae) to extracts of larvae, cocoons and brood food of worker and drone honey bees, Apis mellifera (Hymenoptera: Apidae)

Abstract. Varroa destructor is a parasitic mite of the honey bee species Apis cerana Fabr . and A. mellifera L. Mature females reproduce on the immature stages of their hosts, producing more viable female offspring on drone hosts than on worker hosts. Thus, immature drones are more likely to be infested with mites than immature workers. To investigate the hypothesis that differences in host chemistries underlie the biased distribution of mites between worker and drone brood, the arrestment responses of mites to solvent extracts of a number of stimuli normally encountered by a mite during its life cycle were measured. Mites were arrested by cuticular extracts of worker and drone larvae obtained at 0, 24 and 48 h prior to the time when cell capping is completed. Mites were also arrested by extracts of worker and drone, brood food and cocoons, and by a blend of synthetic fatty acid esters previously shown to be active in the host acquisition process. In a wind tunnel bioassay, mites were attracted to odours from living fifth-instar worker and drone larvae, but not to volatiles from cocoons, brood food or a blend of fatty acid esters. The sex of the host was not an important factor affecting the behavioural responses of the mites in any assay. We conclude that host kairomones play a role in the host acquisition process, but we found no evidence to support the hypothesis that mites use these substances to differentiate between worker and drone brood.     

中草药精油对蜜蜂狄斯瓦螨的熏杀效果

在实验室条件下测定10种中草药精油对蜜蜂狄斯瓦螨Varroa destructor Anderson&Trueman的熏蒸效果,通过测定中草药精油对蜜蜂和螨的毒性,筛选出具有杀螨活性而对蜜蜂较安全的中草药精油。结果表明:中草药精油对蜜蜂都有一定的毒害;而在对蜜蜂无毒性的剂量下,48h时茴香油、丁香油、肉桂油分别能引起92.5%、54%和12%的螨死亡,显著高于对照组(P<0.05);根据初步筛选结果,选择茴香油和丁香油进行进一步毒力测定,结果表明48h时茴香油对螨的LD50为0.949μL,蜜蜂的LD50为4.033μL;丁香油对螨的LD50为0.795μL,蜜蜂的LD50为1.965μL。茴香油在48h时有着较高的选择比(蜜蜂的LD50/螨的LD50=4.250);GC-MS鉴定茴香油最主要成分是茴香脑(占总成分的88.72%),丁香油的主要成分是丁香酚(68.28%)和石竹烯(20.79%)。可见,茴香油和丁香油在控制狄斯瓦螨的应用上具有很大潜力。     

A Comparative Study of Varroa Jacobsoni Reproduction in Worker Cells of Honey Bees (Apis Mellifera) in England and Africanized Bees in Yucatan, Mexico

The aim of this study was to investigate an underlying mechanism of the apparent tolerance of Africanized honey bees (AHB) to Varroa jacobsoni mites in Mexico. This was achieved by conducting the first detailed study into the mites' reproductive biology in AHB worker cells. The data was then compared directly with a similar study previously carried out on European honey bees (EHB) in the UK. A total of 1071 singly infested AHB worker cells were analyzed and compared with the data from 908 singly infested EHB worker cells. There was no significant difference between the number of mother mites dying in the cells (AHB = 2.0%, EHB = 1.8%); the mean number of eggs laid per mite (AHB = 4.86, EHB = 4.93); the number of mites producing no offspring (AHB = 12%, EHB = 9%); and developmental times of the offspring in worker cells of AHB and EHB. However, there was a major difference between the percentage of mother mites producing viable adult female offspring (AHB = 40%, EHB = 75%). This was caused by the increased rate of mite offspring mortality suffered by the first (male) and second (female) offspring in AHB worker cells. Therefore, only an average of 0.7 viable adult female offspring are produced per mite in AHB, compared to 1.0 in EHB.