花蜜微生物与传粉者的相互作用：现状与展望

传粉者是花蜜微生物的重要传播载体,驱动了花蜜微生物群落结构和功能的变化。微生物在花蜜中定殖后,能够改变花蜜质量和花信号,间接影响传粉者觅食决策和适合度,也能通过直接作用影响传粉者健康。本文总结了传粉者对花蜜微生物群落结构和功能的影响,以及花蜜微生物对传粉者觅食行为和适合度的改变,最后阐述了相关领域的未来研究方向,旨在为花蜜微生物和传粉者资源的保护和利用提供参考资料。     

花蜜化学成分及其生态功能研究进展

花蜜化学成分及其生态功能是当今传粉生物学研究中最为活跃的领域之一。花蜜生态功能是基于植物-传粉者的相互作用过程中花蜜各种化学成分生态意义的耦合。目前人们认为花蜜成分是植物为了提高自身繁殖适合度在吸引传粉者、驱避盗蜜者、保护花蜜免受微生物侵染等生态功能上的折中进化。本文综述了国内外有关花蜜中糖类、氨基酸、蛋白质、无机离子、脂类、酚类、生物碱及萜类等化学成分及其生态功能的最新研究结果;总结了花蜜化学成分的变化及其来源、花蜜化学在传粉者类群与传粉综合征中的重要作用;对今后花蜜化学及其生态功能的研究提出了两点建议:一是将分子技术和生态学结合起来,进一步揭示花蜜化学成分形成的分子机理,植物调控花蜜化学成分及其生态适应机制;二是开展群落水平的花蜜化学及其生态功能研究,探讨同期开花植物花蜜化学成分的差异及其与特化、泛化传粉者类群之间的相互关系,确定花蜜化学在群落水平上对植物传粉网络的影响机制。     

常春油麻藤有气味花蜜及其生态功能

一些被子植物能够分泌有气味的花蜜,但这一自然现象很少被关注.作为嗅觉信号线索,有气味的花蜜可能是将访花者和气味信号结合在一起的特征,它与传粉者及盗蜜者的关系值得探索.本研究以常春油麻藤(Mucuna sempervirens)为对象,研究了其开花动态及泊氏长吻松鼠(Dremomys pernyi)和赤腹松鼠(Callosciurus erythraeus)的访花行为,采用顶空固相微萃取和气相色谱-质谱法收集并分析了花蜜的挥发物成分,探讨了花蜜对中华蜜蜂(Apis cerana cerana)的吸引作用及对酸臭蚁(Tapinoma sp.)的毒杀作用.结果表明:常春油麻藤花蜜释放的挥发物以脂肪族化合物为主(87.2％),其中酮类占56.1％,无含硫挥发性成分,这和该属其他蝙蝠传粉种类花蜜释放含硫化合物的结果不一致.此外,常春油麻藤的花蜜对酸臭蚁有慢性毒杀作用,而对中华蜜蜂则有吸引作用.未发现蝙蝠访花,但观察到泊氏长吻松鼠和赤腹松鼠可能为常春油麻藤传粉.因此,常春油麻藤可能不属于蝙蝠传粉的种类.希望本研究能为该属亚洲类群的传粉机制提供数据,并为其他植物类群花蜜成分及功能研究提供新的视角.     

花蜜中酚类物质对群落中同花期植物传粉的影响

显花植物分泌花蜜以奖励传粉昆虫为其提供的传粉服务.许多植物分泌的花蜜中除了糖和氨基酸等物质外,还含有与植物抗虫有关的次生代谢物质,如生物碱和多酚.饲喂试验表明传粉蜂类偏爱采集含有这些次生代谢物质的花蜜.因此,花蜜中的次生代谢物质对分泌这类花蜜的植物传粉具有进化和生态上的意义.但是,花蜜中的次生代谢物质是否会影响群落中其它同花期植物的传粉尚不清楚.在2009年3-5月份,在西双版纳永久样地就中华蜜蜂(Apis cerana)的传粉行为进行研究,比较了两种饲喂条件下,整个蜂群采集花蜜的花粉种类和单个采集蜂携带的花粉团的花粉种类,以及采集蜂蜜囊中花蜜的含糖量(一组饲喂30％蔗糖糖浆,以下简称“喂纯糖浆组”另一组饲喂同样糖浆但含有0.01％槲皮素,以下简称“喂酚糖浆组”).研究结果表明:饲喂两种不同糖浆的蜂群拜访的植物种类数量上没有明显差异.但是,两组采集工蜂所选择采集植物花粉种类却存在明显差异,它们采集的花粉都包含了一些特有的花粉种类.另外,两组采集工蜂所采集含有2种以上花粉种类花粉团的蜜蜂数量存在明显差异,喂酚糖浆组明显高于饲喂纯糖浆组.两组工蜂蜜囊中花蜜的含糖量也存在显著差异,喂酚糖浆组工蜂蜜囊中花蜜的含糖量明显高于喂纯糖浆组.因此,喂酚糖浆组的工蜂拜访植物的专一性比较低,在一次采集过程中倾向拜访花蜜中含糖量高的植物.花蜜中酚类物质增加了群落中同花期植物之间的花粉传播,从而降低了整个群落的传粉效率.     

开花时间对巫山淫羊藿花蜜分泌和结实的影响

运用单因素方差分析研究了巫山淫羊藿(Epimedium wushanense)开花时间对花蜜分泌、有效传粉者活动及结实性的影响.结果表明:(1)开花时间对巫山淫羊藿花蜜常备量和含糖量的影响极其显著.3个种群早期花蜜常备量较高,而花蜜含糖量在晚期较高.(2)开花时间对有效传粉者行为也具显著影响.一天中,有效传粉者的访花活动主要集中在中午;在整个花期进程中,晚期有效访花频率明显大于早期.(3)开花时间对巫山淫羊藿结实率、果实种子数及果实种子败育率的影响均显著,其中早期和中期的结实率和果实种子数大于晚期;而果实种子败育率则在晚期较高.因此,人工栽培巫山淫羊藿应注意花期调控,并选择早期果实进行实验或育种,其效果更好.     

植物微生物组生态功能与群落构建过程研究进展

植物各个器官表面及内部定殖着高度多样化的微生物群落,这些微生物与植物长期共进化,作为宿主植物的“共生功能体”(holobiont)在植物生长发育、养分吸收、病害抵御和环境胁迫适应性等方面发挥了重要作用。得益于近10年来多组学技术的发展和应用,有关植物微生物群落的多样性、组成和功能特征、群落构建的驱动因素和植物–微生物互作机制等方面研究取得了一系列重要进展。然而,与土壤微生物组相比,目前对植物微生物组的认识及其应用尚且不足。本文系统总结了植物微生物组的组成特征,植物微生物在调节植物生长发育、促进养分吸收、提高病害抵御能力及环境胁迫适应性等方面的功能及作用机制,从宿主选择、环境因子以及生物互作3个方面总结了驱动植物微生物群落构建的因素,并着重阐述了植物–微生物互作如何塑造植物微生物群落以及如何调节对植物的有益功能。此外,我们对未来植物微生物组研究和应用面临的挑战进行了展望,如核心微生物组挖掘和合成群落构建,植物–微生物互作的分子调控机制,植物微生物群落水平上的互作机制等。深入理解植物微生物群落特征、生态功能以及构建过程对于精准调控植物微生物组以提高植物适应性和生产力以及维持生态系统健康具有...     

两种生境下野慈姑繁殖差异及其机制

植物的生长环境不但可直接作用于其可利用资源的多寡及其繁殖分配,而且可通过影响植物所在的群落以及传粉者的组成或行为而间接的导致繁殖差异。然而,直接与间接作用的方向或强度很少被同时关注,从而限制了从机制上理解环境对植物繁殖的影响。选取野慈姑(Sagittaria trifolia L.)为研究材料,将相同基因型组成的植株分别种植在光照差异明显的两个同质园(林荫区与日照区),于盛花期对野慈姑的开花数量、昆虫访花进行了观察,确定其主要访花昆虫、记录访花行为以及野慈姑的繁殖产出水平。研究表明,日照区野慈姑每天开放的花朵和植株数量显著高于林荫区。两个区域内野慈姑的主要访花者有四大类:蚜蝇类、蜂类、蝶类和蝇类;其中林荫区的主要传粉昆虫为蚜蝇类,而日照区为蜂类。日照区昆虫单位时间内的访花频率、昆虫每回合访问的雄花数、总花数和花序数均显著高于林荫区。日照区野慈姑的座果率显著高于林荫区,而单果种子数量与种子面积与林荫区相当。总体而言,日照区野慈姑的繁殖产出达到了林荫区的三倍以上,这是环境的直接作用与环境介导的传粉作用在同一方向上叠加的结果,且前者占主导地位。在关注植物、物理环境和生物因子三者之间互作的前提下,量化每个繁殖阶段的水平,特别是传粉昆虫在不同生境下的表现,阐述了环境异质性导致植物繁殖差异的根本原因。     

青藏高原微生物生态功能研究进展

青藏高原作为地球“第三极”和全球气候变化敏感区,有着复杂和极端的气候环境,同时也蕴藏了丰富而特殊的生物资源,因此其生物多样性及其对全球气候变化的响应等研究备受关注。随着免培养的分子生态学和高通量测序等技术的迅速发展,青藏高原微生物的多样性及其功能、与其他生物互作等方面的研究不断深入,人们得以进一步认识青藏高原微生物在调节和维持生态系统平衡稳定等方面的重要作用。本文以青藏高原微生物合成功能中的固碳、固氮和藏区特色食品生产和分解作用中的碳、氮、磷、硫元素循环和有毒有害物质的降解以及共生作用中的与微生物、动物、植物共生为主要内容,讨论了微生物对青藏高原其他物种以及生态环境等方面的独特贡献,并对其生态功能进行总结,明确青藏高原微生物对于其脆弱生态环境的重要意义,以期为青藏高原生物多样性保护及微生物资源的开发利用提供参考。     

植物微生物组的功能及应用研究进展

健康的植物中生活着多种多样但分类学结构不同的微生物群落,它们在所有可接触到的植物组织中定殖.这些微生物群落赋予植物宿主健康优势,包括促进宿主植物生长、营养吸收、抗逆性和对病原菌的抵抗力等.植物菌群及其相互作用具有高度的多样性,多种因素决定着群落的组成和功能.虽然从19世纪开始植物菌群就被人们所认识,但对其功能及应用的相...     

重金属污染土壤植物修复中的微生物功能研究进展

综述了国内外在重金属污染土壤植物-微生物联合修复领域的研究报道,总结了近5年的研究实例。植物-微生物联合修复体系具有生物固定与生物去除土壤重金属的两种功能,根际微生物可以菌根、内生菌等方式与根系形成联合体,通过增强植物抗性和优化根际环境,促进根系发展,增强植物吸收和向上转运重金属的能力。建立植物-微生物联合修复体系,可充分发挥植物与微生物作用功能的优势,提高污染土壤的修复效率。增强植物修复体系中微生物功能的重点是深入研究根际微生物、根系和介质载体三者之间复合功能,结合污染土壤类型与植物群落配置的特点筛选扩繁高效菌种与菌群。     

Effects of nectar-robbing on plant reproduction and evolution

The relationship between plant and pollinator is considered as the mutualism because plant benefits from the pollinator's transport of male gametes and pollinator benefits from plant's reward.Nectar robbers are frequently described as cheaters in the plant-pollinator mutualism,because it is assumed that they obtain a reward (nectar) without providing a service (pollination).Nectar robbers are birds,insects,or other flower visitors that remove nectar from flowers through a hole pierced or bitten in the corolla.Nectar robbing represents a complex relationship between animals and plants.Whether plants benefit from the relationship is always a controversial issue in earlier studies.This paper is a review of the recent literatures on nectar robbing and attempts to acquire an expanded understanding of the ecological and evolutionary roles that robbers play.Understanding the effects of nectar robbers on the plants that they visited and other flower visitors is especially important when one considers the high rates of robbing that a plant population may experience and the high percentage of all flower visitors that nectar robbers make to some species.There are two standpoints in explaining why animals forage on flowers and steal nectar in an illegitimate behavior.One is that animals can only get food in illegitimate way because of the mismatch of the morphologies of animals'mouthparts and floral structure.The other point of view argues that nectar robbing is a relatively more efficient,thus more energy-saving way for animals to get nectar from flowers.This is probably associated with the difficulty of changing attitudes that have been held for a long time.In the case of positive effect,the bodies of nectar robbers frequently touch the sex organs of plants during their visiting to the flowers and causing pollination.The neutral effect,nectar robbers' behavior may destruct the corollas of flowers,but they neither touch the sex organs nor destroy the ovules.Their behavior does not affect the fruit sets or seed sets of the hosting plant.Besides the direct impacts on plants,nectar robbers may also have an indirect effect on the behavior of the legitimate pollinators.Under some circumstances,the change in pollinator behavior could result in improved reproductive fitness of plants through increased pollen flow and out-crossing.     

Effects of nectar-robbing on plant reproduction and evolution

The relationship between plant and pollinator is considered as the mutualism because plant benefits from the pollinator’s transport of male gametes and pollinator benefits from plant’s reward. Nectar robbers are frequently described as cheaters in the plant-pollinator mutualism, because it is assumed that they obtain a reward (nectar) without providing a service (pollination). Nectar robbers are birds, insects, or other flower visitors that remove nectar from flowers through a hole pierced or bitten in the corolla. Nectar robbing represents a complex relationship between animals and plants. Whether plants benefit from the relationship is always a controversial issue in earlier studies. This paper is a review of the recent literatures on nectar robbing and attempts to acquire an expanded understanding of the ecological and evolutionary roles that robbers play. Understanding the effects of nectar robbers on the plants that they visited and other flower visitors is especially important when one considers the high rates of robbing that a plant population may experience and the high percentage of all flower visitors that nectar robbers make to some species. There are two standpoints in explaining why animals forage on flowers and steal nectar in an illegitimate behavior. One is that animals can only get food in illegitimate way because of the mismatch of the morphologies of animals’ mouthparts and floral structure. The other point of view argues that nectar robbing is a relatively more efficient, thus more energy-saving way for animals to get nectar from flowers. This is probably associated with the difficulty of changing attitudes that have been held for a long time. In the case of positive effect, the bodies of nectar robbers frequently touch the sex organs of plants during their visiting to the flowers and causing pollination. The neutral effect, nectar robbers’ behavior may destruct the corollas of flowers, but they neither touch the sex organs nor destroy the ovules. Their behavior does not affect the fruit sets or seed sets of the hosting plant. Besides the direct impacts on plants, nectar robbers may also have an indirect effect on the behavior of the legitimate pollinators. Under some circumstances, the change in pollinator behavior could result in improved reproductive fitness of plants through increased pollen flow and out-crossing. __________ Translated from Acta phytoecologiaca Sinica, 2006, 30(4): 695–702 [译自: 植物生态学报]     

熊蜂盗蜜的花色偏好及对长距忍冬繁殖适合度的影响

植物开花过程中花色的变化作为一种指示信号会引导传粉者偏好访问变色前的花, 那么花色的变化能否影响昆虫盗蜜, 目前还不清楚。为了探究盗蜜者是否偏好变色前的花以及盗蜜对植物繁殖适合度的作用, 我们以忍冬属长距忍冬(Lonicera calcarata)为研究对象, 其花期3天, 花色从白变为黄, 再变为橘红, 野外记录访花昆虫的访问行为, 计算合法访问和盗蜜的频率; 统计不同居群3种颜色花的盗蜜率, 测量它们正常和盗蜜花朵的形态特征, 测量开花中正常和盗蜜花朵的花蜜现存量和糖浓度, 统计正常和盗蜜的花苞中花粉和胚珠总数, 进行不同人工授粉处理统计结实率。长距忍冬的访花者有尺蛾、蝶和熊蜂, 其中熊蜂可以合法地访花和对不同颜色的花盗蜜, 总的盗蜜频率高于合法访花频率; 橘红色和黄色花的盗蜜率、盗洞大小显著高于白色花; 盗蜜花的形态特征(尤其是唇瓣和花药)整体上小于正常花, 盗蜜减少单花的花蜜分泌量, 但不影响其糖浓度; 盗蜜不影响花苞中的花粉总数、胚珠数和植株的结实率。该研究表明盗蜜者熊蜂没有偏好访问变色前的花, 盗蜜会减少花蜜分泌量, 没有直接显著影响植物的雌雄适合度。植物花色变化对访花昆虫行为的影响需要更深入和持续的研究。     

Floral signposts: testing the significance of visual 'nectar guides' for pollinator behaviour and plant fitness

Nectar guides, contrasting patterns on flowers that supposedly direct pollinators towards a concealed nectar reward, are taxonomically widespread. However, there have been few studies of their functional significance and effects on plant fitness. Most previous studies focused on pollinator behaviour and used artificial flowers in laboratory settings. We experimentally investigated the role of putative nectar guides in a natural system: the South African iris Lapeirousia oreogena, whose flowers have a clearly visible pattern of six white arrow-markings pointing towards the narrow entrance of the long corolla tube, and its sole pollinator, a long-proboscid nemestrinid fly. We painted over none, some or all of the white arrow-markings with ink that matched the colour of the corolla background. Although arrow-marking removal had little effect on the approaches by flies to flowers from a distance, it dramatically reduced the likelihood of proboscis insertion. Export of pollen dye analogue (an estimate of male fitness) was reduced to almost zero in flowers from which all nectar guides had been removed, and fruit set (a measure of female fitness) was also significantly reduced. Our results confirm that the markings on L. oreogena flowers serve as nectar guides and suggest that they are under strong selective maintenance through both male and female fitness components in this pollination system.     

大和切叶蜂对其蜜源植物披针叶黄华的盗蜜行为

【目的】调查和观测内蒙古毛乌素沙地大和切叶蜂Megachile (Xanthosaurus) japonica Alfken对其蜜源植物披针叶黄华Thermopsis lupinoides (L.)的盗蜜行为。【方法】在披针叶黄华花期内, 设置样方观测披针叶黄华的主要访花昆虫。采用目测, 拍照等方法对大和切叶蜂盗蜜行为进行观测, 记录和统计花被盗蜜后留下的盗蜜孔的数量和在花上的位置。【结果】大和切叶蜂在披针叶黄华传粉蜂中数量上占有绝对的优势。作为初级盗蜜者时, 用上颚在花基部切割出一个纵向裂口, 将口器伸入孔内吸取花蜜。作为次级盗蜜者时, 利用已有的孔洞来吸蜜。在盗蜜时没有表现出寻找已经存在的盗蜜孔来吸蜜的现象, 同时其个体在盗蜜时表现出“偏好”花基部一侧的行为。在13个样地, 已开放花朵被盗蜜率最低为95.4%, 最高达到100%, 而未开放花朵的被盗蜜率最高则达到64.7%。【结论】在毛乌素沙地大和切叶蜂既是披针叶黄华的主要传粉者, 也是其初级盗蜜者和次级盗蜜者。