蜜蜂级型分化机理

蜜蜂Apis spp.能有效地为多种植物及农作物授粉, 具有重要的经济和生态价值; 蜜蜂作为高度真社会性昆虫, 已成为社会生物学研究的模式生物。社会性昆虫的生殖劳动分工具有重要的进化意义, 而级型分化是形成生殖劳动分工的基础。近年来, 关于蜜蜂级型分化的研究已取得诸多重要成果, 其机理也得到了较为深入的阐释。营养差异引发蜜蜂幼虫的级型分化。蜂王浆中的主要蛋白组分之一--Royalactin是诱导蜂王发育的关键营养因子, 而脂肪体细胞的表皮生长因子受体介导了Royalactin的这种蜂王诱导作用。DNA甲基化是重要的表观遗传机制之一, 且与个体发育和疾病发生紧密相关, 近来的研究表明DNA甲基化在蜜蜂级型分化过程中发挥重要的调控作用。此外, 越来越多的研究进一步深化了人们对内分泌系统调节级型分化作用的认识。本文从关键营养因子调控、 表观遗传调控和内分泌调节3方面综述蜜蜂级型分化的机理, 并对未来的研究提出可能的方向。     

表观遗传学与蜜蜂级型分化的研究进展

蜜蜂是一种高度社会化的昆虫,一个完整健康的蜂群通常是由蜂王、工蜂和雄蜂组成。尽管蜂王和工蜂的遗传物质相同,但它们在形态特征、行为职能和寿命方面表现出显著的差异。许多研究结果表明,营养因素是造成蜜蜂级型分化现象的主要原因,它可以影响蜂王幼虫和工蜂幼虫体内大量基因和蛋白的差异表达。随着表观遗传学的发展,人们对基因表达的调控机制有了新的认识,它与DNA甲基化、非编码RNA调控和组蛋白乙酰化等密切相关。这也为蜜蜂级型分化的分子机制提供了新的理论。本文就表观遗传学和蜜蜂级型分化的研究进展做一综述。     

蜜蜂蜂王与工蜂级型分化研究进展

蜜蜂ApismekiferaL .是典型的社会性昆虫 ,蜂王和工蜂都是由受精卵发育而来的二倍体成蜂 ,但是在形态、生理、行为等方面有明显的差异 ,属于不同的级型。蜂王和工蜂的级型分化的关键时期发生在幼虫的 4龄末至 5龄止。分化是由分化基因调控的 ,幼虫期食物的质和量是分化的外部决定因子。JH对两级型中卵巢的分化有非常重要的调控作用。蜜蜂脑或其它组织中可能有分泌调控CA的咽侧体调节激素 ,它们通过对CA中JH的合成和分泌的调控而参与了分化的调控。章鱼胺等生物胺也参与了分化调控过程。     

LncRNA在蜜蜂级型分化中的功能研究

蜜蜂的级型分化被证实是由蜂王浆中的Royalactin决定,工蜂和蜂王幼虫在级型分化时编码基因的表达差异也被广泛研究.我们发现,在蜜蜂幼虫的级型分化过程中,lncRNA也有着显著的表达差异,因此认为,lncRNA也参与了蜜蜂的级型分化过程.进一步的分析显示,lncRNA可能通过影响上下游基因的转录和功能执行的方式,在蜜蜂早期发育的多细胞组织发育、神经系统发育和转录调控的过程中起到重要的调控作用.     

熊蜂级型分化研究进展

熊蜂是众多野生植物及农作物的有效授粉昆虫,具有重要的经济和生态价值。熊蜂复杂的生长发育过程及社会性使其成为社会生物学研究的最佳模式生物之一。社会性昆虫的生殖劳动分工具有重要的进化意义,而级型分化是形成生殖劳动分工的基础。蜜蜂级型分化的研究已取得诸多重要成果,其机理也得到了较为深入的阐释,而熊蜂的社会性研究尚未形成系统,与蜜蜂研究相差甚远。近来的研究表明,饲喂频率或者饲喂总量的差异能够引起熊蜂级型分化的发生。保幼激素和蜕皮激素与熊蜂幼虫的发育紧密联系,在熊蜂级型分化的过程中发挥重要作用。一些参与蜜蜂级型分化的基因,在熊蜂级型间也存在差异表达。此外,群体间的相互作用以及蜂王和工蜂间的竞争也是促进熊蜂级型分化发生的重要因素。本文从营养、激素调控、群体发展及相互作用等方面综述熊蜂级型分化机制,并对未来的研究提出可能方向。     

蜜蜂级型分化相关生理因子研究进展

蜜蜂Apis melliferaL.是典型的社会性昆虫,蜂群内蜂王和工蜂的级型分化现象吸引了众多研究者的高度关注。蜜蜂级型分化是一个极为复杂的生长发育调节过程,许多重要生理因子参与其中,包括保幼激素、蜕皮激素、胰岛素/胰岛素类似物信号通路等。本文概述了蜜蜂级型分化相关生理因子研究进展:保幼激素决定着蜜蜂级型发育的轨迹并能阻止卵巢发育过程中细胞的程序性死亡;蜕皮激素协同保幼激素发挥调节作用;蜜蜂蜂王与工蜂的躯体大小及相关器官大小的级型特异性差异与胰岛素/胰岛素类似物信号通路密切相关。此外,本文还介绍了蜜蜂级型分化模型,该模型系统描述了上述各生理因子对级型分化的综合作用。     

褐飞虱翊型分化遗传规律的研究

以褐飞虱Nilaparvata lugens (Stal)长翅型(macroptery,简称M)、短翅型(brachyptery,简称B)的遗传纯系为实验材料,进行亲本、稻株生育期、虫口密度三因子交互实验。结果表明：(1)在环境条件(指稻株生育期、虫口密度等)一致时,B♀×B♂、B♀×M♂、M♀×B♂、M♀×M♂四种亲本组合的F1代短翅型成虫百分率分别为：98%、92%、64%、29%,各亲本组合间差异极显著;(2)亲本相同时,将F1代褐飞虱初孵若虫多头词养(多于l0头/株)在黄熟期稻株上,其长翅型雌、雄成虫百分数均高于灌浆期稻株上;(3)单头饲养实验中发现,不论亲本组合、稻株生育期如何,雌虫绝大多数分化为短翅型,而雄虫则几乎全为长翅型。这表明褐飞虱的翅型分化遗传由一个受多种因子影响的调控体系决定,且调控作用与性别有关。     

黄瓜性型分化的分子机制

黄瓜(Cucumis sativus)是雌雄异花植物性型分化研究的重要模式植物,近年来虽然其性型分化的分子机制研究取得了一定的成果,但其性型分化的调控机制尚未完全阐明。该文综合花器官发育基因、性别决定基因、内源激素、环境因子、性型分化假说,在分子水平构建了黄瓜性型分化的表达调控网络。同时对激素和性别决定基因协控的黄瓜单性花器官凋亡机制进行了阐述,并就miRNA在黄瓜性型分化调控中的作用进行了探讨。     

华东地区中华蜜蜂六地理种群的遗传多样性及遗传分化

目的】利用23对微卫星标记对来自于南昌、黄山、桐庐、费县、宜兴、武夷山6个华东地区的中华蜜蜂Apis cerana cerana种群进行遗传多样性及遗传分化分析。【方法】通过计算多态信息含量、平均杂合度、等位基因数、遗传距离、基因流、F 统计量等参数, 评估各中蜂种群遗传多样性和各种群间遗传分化。【结果】各座位的等位基因数为5（A014）至30（AP043）。所有种群均显示较高水平的期望杂合度, 其中, 武夷山中蜂最低, 为0.4280; 南昌中蜂最高, 为0.6329。各中蜂种群间存在极显著的遗传分化, 平均分化系数(Fst)为0.344。基于Nei氏遗传距离运用NJ聚类法将6个中蜂种群划分为3类。【结论】华东6个中蜂种群的遗传多样性较高, 遗传分化显著; 分析遗传分化与地理距离的关系发现, 华东6个中蜂种群间的遗传分化与地理距离不存在显著相关。     

中华蜜蜂种群遗传分化与环境适应机制研究进展

中华蜜蜂Apis cerana cerana是我国十分关键和重要的传粉媒介昆虫,在不同栖息地中广泛分布,表现出飞行敏捷、采蜜期长和适应性强等优点。但近些年中华蜜蜂正面临前所未有的种群多样性下降,为保护这些特定的中华蜜蜂种群的遗传资源,研究人员基于几何形态学、分子生物学和基因组学等技术手段对中华蜜蜂遗传分化和环境适应机制进行了研究,而在我国这些多样化的中华蜜蜂种群为揭示其种群适应性进化规律提供了绝佳材料。本文从中华蜜蜂种群遗传分化与环境变化的相关性、形态变异与环境适应、环境适应性生理和行为改变及这些表型变化的遗传机制4个方面综述了相关研究进展。研究发现,物理屏障和生态环境的变化,特别是海拔和纬度相关环境变化是造成中华蜜蜂种群分化的主要原因。气候因子中,温度、氧气、辐射和湿度等对中华蜜蜂环境适应性的形态发育和生理生态特征产生重要影响。形态特征的变化主要体现在体型大小和体色上,代谢生理和行为改变是其环境适应的重要策略。基于现代基因组学和分子生物学技术的种群遗传学和种群基因组学研究发现,与社会分工行为、学习感知行为、信息感知、生长发育、温度适应和代谢等相关的基因或通路受到自然选择作用,这为中华...     

mRNA expression and DNA methylation in three key genes involved in caste differentiation in female honeybees(Apis mellifera)

In honeybee(Apis mellifera)colonies,queens and workers are alternative forms of the adult female honeybee that develop from genetically identical zygotes but that depend on differential nourishment.Queens and workers display distinct morphologies,anatomies and behavior,better known as caste differentiation.Despite some basic insights,the exact mechanism responsible for this phenomenon,especially at the molecular level,remains unclear although some progress has been achieved.In this study,we examined mRNA levels of the TOR(target of rapamycin)and Dnmt3(DNA methyltransferase 3)genes,closely related to caste differentiation in honeybees.We also investigated mRNA expression of the S6K(similar to RPS6-p70-protein kinase)gene linked closely to organismal growth and development in queen and worker larvae(1-day and 3-day old).Last,we investigated the methylation status of these three genes in corresponding castes.We found no difference in mRNA expression for the three genes between 1st instar queen and worker larvae;however,3rd instar queen larvae had a higher level of TOR mRNA than worker larvae.Methylation levels of all three genes were lower in queen larvae than worker larvae but the differences were not statistically significant.These findings provide basic data for broadening our understanding of caste differentiation in female honeybees.     

Nutritional versus genetic correlates of caste differentiation in a desert ant

1. In many ant species, caste differentiation stems from trophic differences at the larval stage. Adult workers that feed larvae have great control over the allocation of colony resources to growth (production of workers) versus reproduction (production of queens). However, larval caste fate may also be constrained very early on through direct genetic effects or non‐genetic maternal effects. 2. Here, we combined isotopic and genetic analyses to study the developmental origin of queens and workers in a desert‐dwelling ant, Cataglyphis tartessica (Amor & Ortega, 2014). Queens do not found new colonies alone but rather disperse with workers. As the latter are always wingless, selection pressures on specific queen traits such as flight ability have become relaxed. Though the phylogenetically related species, C. emmae (Forel, 1909) only produces winged queens much larger than workers, C. tartessica produces two types of small queens relative to workers: brachypterous (short‐winged) queens and permanently apterous ergatoid (wingless and worker‐like) queens. 3. Upon emergence, workers and ergatoids have similar δ15N isotopic values, which were lower than those of brachypters, suggesting the latter are fed more protein as larvae. Microsatellite analyses indicated that: (i) colonies are mostly monogynous and monandrous; (ii) both ergatoids and brachypters are equally related to workers; and (iii) in the few polyandrous colonies, patrilines were evenly represented across workers, brachypters and ergatoids. 4. Overall, there was no evidence of genetic caste determination. We suggest that, in contrast to brachypters, ergatoids are selfish individuals that escape the nutritional castration carried out by workers and develop into queens in spite of the colony's collective interests.     

Juvenile hormone titers and caste differentiation in the damp-wood termite Hodotermopsis sjostedti (Isoptera, Termopsidae)

Termites are social insects, presenting morphologically distinct castes, performing specific tasks in the colony. The developmental processes underlying caste differentiation are mainly controlled by juvenile hormone (JH). Although many fragmentary data support this fact, there was no comparative work on JH titers during the caste differentiation processes. In this study, JH titer variation was investigated using a liquid chromatography-mass spectrometry (LC-MS) quantification method in all castes of the Japanese damp-wood termite Hodotermopsis sjostedti, especially focusing on the soldier caste differentiation pathway, which was induced by treatment with a JH analog. Hemolymph JH titers fluctuated between 20 and 720pg/microl. A peak of JH was observed during molting events for the pseudergate stationary molt and presoldier differentiation, but this peak was absent prior to the imaginal molt. Soldier caste differentiation was generally associated with high JH titers and nymph to alate differentiation with low JH titers. However, JH titer rose in females during alate maturation, probably in relation to vitellogenesis. In comparison, JH titer was surprisingly low in neotenics. On the basis of these results in both natural and artificial conditions, the current model for JH action on termite caste differentiation is discussed and re-appraised.     

社会性昆虫级型和行为分化机制研究进展

社会性的出现是生物演化过程中的重要革新, 理解社会性的演化和调控机制具有重要的理论和实际意义。社会性昆虫的个体间有着明显的级型分化和劳动分工, 这有利于它们适应复杂的环境变化。理解社会性昆虫如何产生不同的形态、行为和生活史特性, 一直是进化和发育生物学的重要目标。随着测序技术的不断更新及生物信息学的快速发展, 已经有众多关于社会性昆虫级型和行为分化机制的研究报道。本文通过整理社会性昆虫研究的已有成果, 从环境因素、生理调控和分子机制等方面对社会性昆虫级型和行为分化机制相关研究进展进行了综述, 并对未来的研究方向做出了展望。根据现有证据, 社会性昆虫所生活的生物环境(食物营养、信息素、表皮碳氢化合物)和非生物环境(温度、气候等)均能直接或间接影响社会性昆虫级型和行为的分化; 保幼激素、蜕皮激素、类胰岛素及生物胺等内分泌激素和神经激素对社会性昆虫的级型和行为分化也有重要的调控作用; 此外, 遗传因素、新基因等DNA序列或基因组结构上的变化以及表观遗传修饰、基因的差异表达等基因调控机制均能不同程度地影响社会性昆虫的行为分化。本文建议加强昆虫纲其他社会性类群如半翅目蚜虫和缨翅目蓟马等的社会性行为及其演化机制的研究, 以加深对社会性昆虫起源及其行为演化的理解和认识。     

Development and evolution of caste dimorphism in honeybees – a modeling approach

The difference in phenotypes of queens and workers is a hallmark of the highly eusocial insects. The caste dimorphism is often described as a switch‐controlled polyphenism, in which environmental conditions decide an individual's caste. Using theoretical modeling and empirical data from honeybees, we show that there is no discrete larval developmental switch. Instead, a combination of larval developmental plasticity and nurse worker feeding behavior make up a colony‐level social and physiological system that regulates development and produces the caste dimorphism. Discrete queen and worker phenotypes are the result of discrete feeding regimes imposed by nurses, whereas a range of experimental feeding regimes produces a continuous range of phenotypes. Worker ovariole numbers are reduced through feeding‐regime‐mediated reduction in juvenile hormone titers, involving reduced sugar in the larval food. Based on the mechanisms identified in our analysis, we propose a scenario of the evolutionary history of honeybee development and feeding regimes.     

社会性昆虫品级分化的基因调控

品级分化是社会性昆虫的主要特征之一,包括雌性分化为生殖个体和非生殖个体,这一分化过程是受基因调控的。目前对社会性昆虫品级分化相关基因表达研究的报道较多,但是对这一分化的调控机制并不十分清楚,对这些基因的来源还存在争议。文章对社会性昆虫品级分化相关基因的研究概况作一综述,以期对以后相关方面的深入研究有所帮助。     

Soldier caste influences on candidate primer pheromone levels and juvenile hormone-dependent caste differentiation in workers of the termite Reticulitermes flavipes

Caste systems and the division of labor they make possible are common underlying features of all social insects. Multiple extrinsic factors have been shown to impact caste composition in social insect colonies. Primer pheromones are one type of extrinsic caste-regulatory factor; they are chemical signaling molecules produced by certain colony members to impact developmental physiology of recipient nestmates. However, only limited evidence exists regarding primer pheromones and their actions in eusocial termites. In previous research we identified two soldier-produced terpenes, γ-cadinene (CAD) and γ-cadinenal (ALD), as candidate primer pheromones of the lower termite Reticulitermes flavipes. In the present study we tested hypotheses related to CAD and ALD action in recipient individuals. We examined the influences of terminally developed soldier termites on (1) CAD and ALD levels and (2) caste differentiation in developmentally totipotent workers. Our findings show CAD and ALD (respectively) are caste stimulatory and inhibitory components of chemical blends present in soldier heads, ALD levels increase significantly (10.9×) in workers only in the presence of soldiers, and soldiers can reduce developmental-hormone response thresholds of workers, presumably via ALD action. These findings provide novel evidence supporting that CAD and ALD are authentic caste-regulatory primer pheromones in Reticulitermes.