蜜蜂无政府主义蜂群的研究进展

西方蜜蜂Apis mellifera作为典型的社会性昆虫, 最重要的特征是生殖劳动分工。蜂王垄断蜂群的生殖权利, 工蜂生殖功能受到抑制, 从事除产卵和交配以外的所有职能。而在无政府主义蜂群中, 即使蜂王存在, 也有较多工蜂的卵巢激活并产卵, 蜂群中大多数雄蜂是工蜂的后代。这些特殊蜂群为正常蜂群工蜂不育机制研究提供了绝佳的反例材料。本文对无政府主义蜂群的行为特征、 产生条件、 遗传基础等研究进行了综述。无政府主义蜂群中有较多的工蜂产卵, 且工蜂所产卵能够逃避工蜂监督, 这种行为的产生受环境、 遗传组成、 基因表达等多种因素的影响, 并且遗传结构体系复杂, 参与调控的基因数量多。无政府主义蜂群行为机制的研究为工蜂不育机制的揭示及其他社会性昆虫工职不育基因的筛选和功能研究提供借鉴。     

蜜蜂群内生殖分工体系的形成及其维持

本文对蜜蜂群内生殖分工体系的形成及其维持机制进行综述。蜜蜂群体具有完善的劳动分工(包括生殖分工)体系,蜂王垄断生殖权力,而工蜂生殖器官发育不完全,在蜂王信息素和幼虫信息素的作用下产卵受到抑制。蜂王的多雄交配机制降低了群内个体间的亲缘关系,但也促进了工蜂间相互监督机制的形成。工蜂间的相互监督,结合蜂王和幼虫信息素对工蜂卵巢发育的影响,解决了蜂王与工蜂、工蜂与工蜂间的生殖利益冲突,保障了蜂群内的生殖分工体系,提高了群体效率,维护了蜂群的真社会性。     

意大利工蜂前胃显微结构观察

通过解剖镜和扫描电子显微镜对意大利工蜂Apismellifera ligusticaL.前胃的形态结构进行了观察,发现前胃包括前胃瓣和前胃管两部分。前胃瓣是由4个瓣片环围而成,球形,在蜜囊中,每个瓣片的长度约为0.8mm,前端为三角形结构,可以运动,边长约为0.35mm,三角形的前缘有较长的毛形结构。在每个三角形顶端各有一个直径约为50μm瘤形突起。前胃瓣最大孔径为0.89～1.00mm,具有调整和控制食物进入中肠的功能。前胃管是一直径0.01～0.02mm的较为柔软的管道,插入中肠5mm,该结构可防止中肠食物的倒流。前胃瓣上有7种毛状结构,分布在前胃瓣的不同部位,该结构的功能是分离和过滤、捕捉和感觉功能。本研究为理解工蜂前胃在消化系统中的功能和作用以及昆虫形态学和昆虫分类学提供理论依据。     

生物行为变化研究的新模式——工蜂

蜜蜂群体由1只蜂王、几百只雄蜂和数千只工蜂组成,工蜂数量巨大,除蜂王和雄蜂共同完成生殖任务,巢内外活动均由工蜂完成,其行为呈现多样性。工蜂发育经过卵、幼虫、蛹、成虫4个阶段,其活动范围由温度、湿度相对稳定的巢内环境发展到复杂的巢外自然环境。随发育阶段和生活环境的变化,工蜂的生理、行为等方面也发生相应变化,这种变化为综合研究生物行为的分子机理提供了可能。又因工蜂数量多、体积较大易于观察、标记、饲养管理简单,而且目前对其形态、发育、生理、分子生物学、神经、社会生态等各方面的研究比较充分,所以工蜂成为研究生物体行为变化、发育机理和个体与群体关系的理想模式生物。该文介绍工蜂活动由巢内发展到巢外时出现的一系列变化以及部分变化的机制,主要包括行为的变化、激素的分泌、代谢活动、飞行能力、神经系统等。     

工蜂利它行为的背后

真社会性的昆虫如蜜蜂的个体间有等级之分，有蜂后、雄蜂和工蜂。其中占绝大多数的工蜂与蜂后一样，有正常的卵巢，都是雌性，而且是蜂后的女儿。她们无私地与同巢伙伴共享食物，共同养育蜂后的后代，但通常不产卵。工蜂为什么不生育？科学家一直在寻找这一问题的答案。起先科学家认为，工蜂可能自愿地选择帮助她们的母亲照顾其他的后代，因为这样比她们自身繁殖传种接代更为有效。然而，最新的研究表明，这类昆虫的真社会并不像以前所说的是一个和谐的“公社”，而相反它是一个微型的专制社会。     

基础知识讲座 蚊虫的遗传防治

(前见本刊1983年第4期及1984年第1期)三、有害基因的利用在此所述的昆虫有害基因是指昆虫群体中存在一种对其本身有害的基因突变,而且有可能在其后代中表现出死亡。这些有害基因的突变可以在群体中自然发生,亦可用物理或化学方法诱发。在研究此类领域时,必须防止所出现的昆虫有     

蚕类昆虫线粒体DNA研究及其在起源与进化研究中的应用

线粒体DNA(mtDNA)属母系遗传,进化速率较核基因快且基因组结构相对简单,已作为理想的分子标记广泛应用于昆虫群体遗传学及分子系统学等研究。本文对蚕类昆虫线粒体DNA在分子水平上的最新研究进展进行了较详细的阐述,重点介绍了蚕类昆虫线粒体基因组的组成及特征、mtDNA克隆与多态性及在蚕类昆虫分子系统学研究中的应用等。     

云南澜沧江流域传粉昆虫——熊蜂多样性现状与保护对策

本文对澜沧江流域内热带地区的勐腊县、亚热带地区的南涧县和寒温带地区的德钦县3个低、中、高海拔地带的重要传粉昆虫——熊蜂属(Bombus)的多样性现状进行了观察研究,并与20年前的资料进行了比较分析。结果得出：20多年以来,由于植被人为的破坏严重,传粉昆虫的自然生存环境发生了较大的改变,致使3个不同地带的传粉昆虫一部分物种消失,一部分物种的群体减少,而少数原来极少采到或者未采到的物种和种群则成为优势群体。从而得出：在云南省澜沧江流域人为的生境改变,对传粉昆虫的物种多样性影响极大,要保护或者恢复传粉昆虫的种群,首要任务是保护和恢复生态系统。     

昆虫行为的左右非对称性研究进展

大脑和行为的左右非对称性一直以来被认为是人类特有的,现在广泛存在于脊椎动物中,而近期又发现无脊椎动物中也存在行为偏侧性。近年来大量研究表明,群居型昆虫的嗅觉行为和大脑的非对称性表现在种群水平上,少数散居型昆虫行为的非对称性往往表现在个体水平上。当前,昆虫行为的单侧性是一个重要的研究领域,有助于对生物多样性中单侧性的早期起源有一个更深刻的认识,对探索昆虫进化发展具有重要的科学意义。对大多数昆虫而言,行为的偏侧性都是通过对触角的左右非对称性研究来实现的,触角作为昆虫感觉系统的重要组成部分,在昆虫寄主定位、识别、取食、觅偶、交配、繁殖、栖息、防御与迁移、稳定飞行速率等社会性行为过程中起着极为重要的作用。本文回顾了国内外膜翅目、双翅目、半翅目、蜚蠊目和直翅目昆虫行为非对称性研究的最新进展,有助于分析不同种类昆虫在进化历史上的相似性以及探讨昆虫在个体水平和群体水平上的进化差异。     

昆虫的标记研究法

在昆虫学技术中,采用标记方法,可以研究昆虫的迁移、越冬、定向、扩散速率、寿命、配偶行为、蜕皮次数、活动范围、寄主更换、传播毒病、捕食现象和寄生现象等一系列的生态学和生物学问题。此外,利用标记方法,还可以推算在一定时间和一定地点的昆虫种群的数量,因此,长时期以来,这种方法,一直受到人们的重视。 标记昆虫的方法很多,但大体上可以归纳为五类,即畸志标记、符签标记、油漆与染料标记、萤光物质标记和放射性同位素标记。在这些标记方法中,畸态标记和符签标记,一次操作,只能使个别昆虫带上标记符号,因此它们只是一种个体标记方法。相反地,如果一次操作,能给大量昆虫带上标记符号,就是群体标记方法。油漆与染料标记、萤光物质标记和放射性同位素标记,既可应用于个别昆虫,又可应用于昆虫的群体,因此,它们既是一种个体标记方法,又是一种群体标记方法。实际上,在现代昆虫学的研究中,有些标记方     

Specialization in policing behaviour among workers in the ant Pachycondyla inversa

Most animal societies are non-clonal and thus subject to conflicts. In social insects, conflict over male production can be resolved by worker policing, i.e. eating of worker-laid eggs (WLE) or aggression towards reproductive workers. All workers in a colony have an interest in policing behaviour being expressed, but there can be asymmetries among workers in performing the actual behaviour. Here, we show that workers of the ant Pachycondyla inversa specialize in policing behaviour. In two types of behavioural assays, workers developed their ovaries and laid eggs. In the first experiment, reproductive workers were introduced into queenright colonies. In the second experiment, WLE were introduced. By observing which individuals policed, we found that aggressive policing was highly skewed among workers that had opportunity to police, and that a similar tendency occurred in egg policing. None of the policing workers had active ovaries, so that policing did not incur a direct selfish benefit to the policer. This suggests that policing is subject to polyethism, just like other tasks in the colony. We discuss several hypotheses on the possible causes of this skew in policing tasks. This is the first non-primate example of specialization in policing tasks without direct selfish interests.     

A test of worker policing theory in an advanced eusocial wasp, Vespula rufa

Mutual policing is an important mechanism for maintaining social harmony in group-living organisms. In some ants, bees, and wasps, workers police male eggs laid by other workers in order to maintain the reproductive primacy of the queen. Kin selection theory predicts that multiple mating by the queen is one factor that can selectively favor worker policing. This is because when the queen is mated to multiple males, workers are more closely related to queen's sons than to the sons of other workers. Here we provide an additional test of worker policing theory in Vespinae wasps. We show that the yellowjacket Vespula rufa is characterized by low mating frequency, and that a significant percentage of the males are workers' sons. This supports theoretical predictions for paternities below 2, and contrasts with other Vespula species, in which paternities are higher and few or no adult males are worker produced, probably due to worker policing, which has been shown in one species, Vespula vulgaris. Behavioral observations support the hypothesis that V. rufa has much reduced worker policing compared to other Vespula. In addition, a significant proportion of worker-laid eggs were policed by the queen.     

Worker reproduction and policing in insect societies: an ESS analysis

Insect societies are vulnerable to exploitation by workers who reproduce selfishly rather than help to rear the queen's offspring. In most species, however, only a small proportion of the workers reproduce. Here, we develop an evolutionarily stable strategy (ESS) model to investigate factors that could explain these observed low levels of reproductive exploitation. Two key factors are identified: relatedness and policing. Relatedness affects the ESS proportion of reproductive workers because laying workers generally work less, leading to greater inclusive fitness costs when within-colony relatedness is higher. The second key factor is policing. In many species, worker-laid eggs are selectively removed or 'policed' by other workers or the queen. We show that policing not only prevents the rearing of worker-laid eggs but can also make it unprofitable for workers to lay eggs in the first place. This can explain why almost no workers reproduce in species with efficient policing, such as honeybees, Apis, and the common wasp, Vespula vulgaris, despite relatively low relatedness caused by multiple mating of the mother queen. Although our analyses focus on social insects, the conclusion that both relatedness and policing can reduce the incentive for cheating applies to other biological systems as well.     

The policing behavior ‘immobilization’ towards ovary-developed workers in the ant, Diacamma sp. from Japan

Summary. In social Hymenoptera worker policing that inhibits direct reproduction of workers occurs mainly in the two ways: (1) destruction of worker progenies (postovipositional policing), and (2) aggression towards ovary-developed workers (preovipositional policing). In the queenless ponerine ant, Diacamma sp. from Japan, the existence of the former type of worker policing has been reported, whereas previous studies have failed to find the latter type. We examine the presence of the latter type of worker policing in this species using more careful observational methods. By a series of experiments reuniting previously separated orphaned and non-orphaned subcolonies we found the following facts. Immediately after the colony reunification, aggression frequently took place. Unlike the one-on-one dominance interaction that occurs in non-manipulated colonies, aggression was often directed towards a single victim from multiple attackers, grasping and pulling the victim. The duration of each aggressive interaction was on average far longer in this situation than that of usual dominance interactions. Most victims consisted of ex-orphan workers, while the majority of the attackers were ex-non-orphan workers. Dissection after the above behavioral observation revealed that the ovaries of the victims were on average more active, often containing mature oocytes, than those of non-victims, while the ovaries of attackers were always inactive. The above findings indicate that worker policing via immobilization, which has been reported in some other Ponerinae, also exists in Diacamma sp. from Japan.Received 22 April 2003; revised 28 June and 19 August 2004; accepted 1 September 2004.     

REPRODUCTIVE CONFLICT IN BUMBLEBEES AND THE EVOLUTION OF WORKER POLICING

Worker policing (mutual repression of reproduction) in the eusocial Hymenoptera represents a leading example of how coercion can facilitate cooperation. The occurrence of worker policing in “primitively” eusocial species with low mating frequencies, which lack relatedness differences conducive to policing, suggests that separate factors may underlie the origin and maintenance of worker policing. We tested this hypothesis by investigating conflict over male parentage in the primitively eusocial, monandrous bumblebee, Bombus terrestris. Using observations, experiments, and microsatellite genotyping, we found that: (a) worker‐ but not queen‐laid male eggs are nearly all eaten (by queens, reproductive, and nonreproductive workers) soon after being laid, so accounting for low observed frequencies of larval and adult worker‐produced males; (b) queen‐ and worker‐laid male eggs have equal viabilities; (c) workers discriminate between queen‐ and worker‐laid eggs using cues on eggs and egg cells that almost certainly originate from queens. The cooccurrence in B. terrestris of these three key elements of “classical” worker policing as found in the highly eusocial, polyandrous honeybees provides novel support for the hypothesis that worker policing can originate in the absence of relatedness differences maintaining it. Worker policing in B. terrestris almost certainly arose via reproductive competition among workers, that is, as “selfish” policing.     

Cape honeybees, Apis mellifera capensis, police worker-laid eggs despite the absence of relatedness benefits

In the Cape honeybee, Apis mellifera capensis, workers lay diploid(female) eggs via thelytoky. In other A. mellifera subspecies,workers lay haploid (male) eggs via arrhenotoky. When thelytokousworker reproduction occurs, worker policing has no relatednessbenefit because workers are equally related to their sisterworkers' clonal offspring and their mother queen's female offspring.We studied worker policing in A. m. capensis and in the arrhenotokousAfrican honeybee A. m. scutellata by quantifying the removalrates of worker-laid and queen-laid eggs. Discriminator coloniesof both subspecies policed worker-laid eggs of both their ownand the other subspecies. The occurrence of worker policing,despite the lack of relatedness benefit, in A. m. capensis stronglysuggests that worker reproduction is costly to the colony andthat policing is maintained because it enhances colony efficiency.In addition, because both subspecies policed each others eggs,it is probable that the mechanism used in thelytokous A. m.capensis to discriminate between queen-laid and worker-laideggs is the same as in arrhenotokous A. m. scutellata.     

Worker reproduction in the ant Formica fusca

We studied the kin conflict over male parentage in the ant Formica fusca. The conflict arises because each worker and queen is most related to her own sons and is thus predicted to lay eggs. Microsatellite analysis of eggs revealed that workers laid eggs in more than half the queenright experimental nests. Nevertheless, almost exclusively diploid offspring were reared in the presence of a queen. This also occurred when worker-laid haploid male eggs were experimentally introduced in to the nests. Because our experimental setup allowed us to exclude the possibility of queen policing, we conclude that worker laid eggs are removed by other workers, either as a response to their parentage or gender. Our results suggest that worker reproduction in F. fusca is ultimately an interplay of conflicts over male parentage and sex allocation and that both worker and self policing have roles as proximate mechanisms of resolution.     

Colony stage and not facultative policing explains pattern of worker reproduction in the Saxon wasp

Inclusive fitness theory predicts that in colonies of social Hymenoptera headed by a multiple‐mated queen, workers should benefit from policing eggs laid by other workers. Foster & Ratnieks provided evidence that in the vespine wasp Dolichovespula saxonica, workers police other workers’ eggs only in colonies headed by a multiple‐mated queen, but not in those headed by a single‐mated one. This conclusion, however, was based on a relatively small sample size, and the original study did not control for possible confounding variables such as the seasonal colony progression of the nests. Our aim, therefore, was to reinvestigate whether or not facultative worker policing occurs in D. saxonica. Remarkably, our data show that in the studied Danish population, there was no correlation between worker–worker relatedness and the percentage of worker‐derived males. In addition, we show that variability in cuticular hydrocarbon profiles among the workers did not significantly correlate with relatedness and that workers therefore probably did not have sufficient information on queen mating frequency from the workers’ cuticular hydrocarbon profiles. Hence, there was no evidence that workers facultatively policed other workers’ eggs in response to queen mating frequency. Nevertheless, our data do show that the seasonal progression of the nest and the location in which the males were reared both explain the patterns of worker reproduction found. Overall, our results suggest that the earlier evidence for facultative worker policing in D. saxonica may have been caused by accidental correlations with certain confounding variables, or, alternatively, that there are large interpopulation differences in the expression of worker policing.     

Worker policing in the common wasp Vespula vulgaris is not aimed at improving colony hygiene

In insect societies, eggs laid by workers are frequently killed by other workers – a behaviour known as “worker policing”. The traditional explanation of worker policing is that it is a mechanism to resolve intracolony conflict, and maintain the reproductive monopoly of the queen. Recently, Pirk et al. (2004) proposed that worker policing instead is aimed at removing unviable worker-laid eggs and is ultimately just another example of hygienic behaviour. Here we test this hypothesis for the common wasp Vespula vulgaris, a species with highly effective worker policing. We show that worker-laid eggs from queenless colonies have a lower hatch rate (68%) than queen-laid eggs (82%). Analysis of egg laying rates of queens and workers, however, shows that the difference is not big enough to explain the apparent absence of adult worker-derived males in this species. Received 30 January 2006; revised 2 May 2006; accepted 5 May 2006.