刺五加、短梗五加的开花动态及繁育系统的比较研究

野外定位观测刺五加（Eleutherococcus senticosus),短梗五回（E.sessiliflorus)的开花进程,花朵的功能形态特征和开花动态,用杂交指数（OCI）,花粉－胚珠比（P／O）,去雄,套袋,人工授粉等方法分别测定刺五加,短梗五加的繁育系统,结果显示,刺五加种群,短梗五加种群的花期均持续1个月左右,刺五加比短梗五加早开花20d 左右,二均有雄蕊先熟现象,刺五加是单全异株植物,种群内既具有雄性,又具有雌株,还具有两性株,繁育系统主要为异交,需要传粉活动才能完成授粉过程,与刺五加不同,短梗五加仅具两性花,但繁育系统也以异交为主,短梗五加两性花中的雌,雄器官既在空间上分离,又在时间上分离,只能进行同株异花间或异株,异花间传粉才能受精结实。     

刺五加传粉生物学研究

本文报道刺五加Eleutherococcus seticosus(Rupr．et Maxim．)Maxim的花朵酬物、访花者类别、访花 者在花序上的访花行为、访花频率及传粉效果。主要结果如下:(1)每朵雄花可提供的报偿是58 000～ 81 000粒花粉,5.5～8.0 µl／d×2～4d花蜜。每朵雌花仅可提供4.0～8.0µl／d× 2～3 d花蜜,不能提 供花粉。每朵两性花可提供19 000～54 000粒花粉和7.0～10.0 µl／d× 2～6 d花蜜。(2)花粉提供者 (雄花和两性花)提供报偿的日期是开花后1～3天,花粉接受者(雌花和两性花在柱头外翻变白时)是 开花后5～7或7～9天,这进一步证实了剌五加雄蕊先熟。另外,花粉提供者提供花蜜的时间在一天中 是8:30至15:30,其高峰是9:00～15:00;花粉接受者提供花蜜的时间在一天中是10:30至16:30,其高 峰是11:00～16:00。这种时间差异可能是刺五加影响传粉者流向(从花粉提供者到花粉接受者)的关键 因素。(3)在刺五加花朵上记录到的访花昆虫有50余种,分别隶属于膜翅目、鳞翅目、鞘翅目、双翅目和 半翅目。在不同天气、不同生境、不同性别的植株记录到的访花频率及高峰时间不同:花粉提供者的 访花高峰早于花粉接受者的;雄株上的访花频率高于雌株的;两性株接受花粉时的访花频率高于提供花 粉时的;而天气越晴朗,刺五加地块(patch)越大,则访花频率越高;每日接受光照越早,访花高峰越早。 (4)从传粉效果看,刺五加种子的形成完全依赖于传粉昆虫的活动。其中,蜂类,如熊蜂、花蜂、切叶蜂、胡蜂和蜜蜂等是最有效的传粉者,其它昆虫如粉蝶、食蚜蝇、寄蝇和鳃角金龟等对传粉也有帮助。     

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.     

栽培细叶百合的传粉生态

对栽培的细叶百合进行传粉生态学的初步观察结果显示：细叶百合2:30～3:00开始开花,5:30～7:00完全开放;单花开放时间3～4 d,群体花期为12 d。花蜜分泌在开花当日,分泌高峰在9:00～11:00,花蜜分泌与昆虫访花谐调一致。花蜜成分主要为葡萄糖、阿拉伯糖和β－D吡喃果糖,花朵挥发物的成分是软脂酸、软脂酸甲酯、肉豆蔻酸、月桂酸、3－叔戊基戊酮、十三烷和1－甲基,2异丙烯基环丁基乙醇。访花昆虫主要是蝶类、蜂类和蝇类;传粉昆虫为绿豹蛱蝶、大红蛱蝶、绢粉蝶及单齿切叶蜂、小齿淡脉隧蜂,其中小齿淡脉隧蜂的传粉活动最为活跃。晴天时小齿淡脉隧蜂访花频率的日变化与光照的日变化呈极显著的正相关,与温度的日变化呈显著的正相关,与湿度的日变化呈显著的负相关;阴天时与空气温度相关极显著,与空气湿度相关显著。细叶百合以异花虫媒传粉为主,部分自交可育。     

花蜜微生物及其生态功能研究进展

花蜜是虫媒植物提供给传粉者最有效的报酬,对花蜜特征介导的植物-传粉者相互关系的研究已成为当今传粉生物学研究中最活跃的领域之一。开花植物分泌的原始花蜜是无菌的,不过一些微生物可经由空气传播至花蜜或(和)通过与传粉者的喙接触而聚集于花蜜中,并利用花蜜中的营养物质进行快速繁殖。花蜜的高渗透压环境导致花蜜中微生物(酵母菌,细菌)的物种多样性相对较低。此外,某些生物(传粉者组成,微生物间的竞争)与非生物因素(渗透压,糖组成,次生代谢物质,抗菌化合物,可利用氮源,温度,pH)也可影响花蜜中微生物群落的形成。花蜜中微生物的代谢活动能够改变花蜜物理(温度,粘度)与化学(pH,H_2O_2含量,糖组成和浓度,氨基酸组分和浓度,以及气味)特性,进而影响传粉者的访花行为与植物的繁殖适合度。因而,对花蜜中微生物及其生态功能的研究近年来颇受传粉生物学家的关注。在总结已发表研究成果的基础上,提出今后的研究有必要结合分子生物学与化学分析技术,以进一步揭示影响花蜜中微生物群落的潜在因素的作用机制,同时对花蜜微生物改变花蜜的物理、化学特性及植物-传粉者之间相互作用的可能原因进行更详尽的阐释,特别是对花蜜微生物在生态系统中所发挥的生态功能进行进一步的研究与认识。     

锦鸡儿(Caragana sinica(Buchoz) Rehd)传粉生物学研究

对唐家河国家自然保护区内锦鸡儿(Caragana sinica(Buchoz)Rehd)的花期、访问者、有效传粉者种类及行为、花蜜分泌节律和繁育系统等进行定点观察和实验处理。结果表明:锦鸡儿单花花期约5d,种群花期约20d。拥有以吸蜜为主的条蜂属(Anthophora)、熊蜂属(Bombus)和太阳鸟属(Aethopyga)3种传粉者功能群。综合访花频率和花粉落置数,茜条蜂(Pyganthophora rubricus Dours)和蓝喉太阳鸟(Aethopyga gouldiae)为主要有效传粉者,熊蜂(Bombus)为次要有效传粉者;蚂蚁(formicidae)和西方蜜蜂(Apis mellifera Linnaeus)为主要盗蜜者。套袋处理结果表明,花蜜分泌日变化中,最大花蜜量发生在16:00左右;花蜜随单花花期变化中,花蜜体积最大发生于单花第5天;分泌均呈递增趋势。花蜜分泌节律与访花频率呈正相关。繁育系统检测显示,花粉胚珠比为3779±320.92,柱头最适可授期在开花第2—3天;结合不同授粉处理表明:锦鸡儿自交不亲和,需要传粉者,为蜂媒兼鸟媒植物。     

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.     

鹅掌楸的传粉环境与性配置

近年来研究表明,动物传粉者对植物花部诱物特征（花冠形状大小,花蜜产量和花序大小）有潜在的选择作用。不同效率的传粉者可能导致植物性配置的变化,不同传粉效率的环境下,两性花植物鹅掌楸的花粉和胚珠的配置不同,居群Ｚ在主要访花者是传粉效率较低的类的传粉环境下,花粉粒小,花粉数量相对较高;另一方面胚珠投资的减少,缓解了低效的传粉（增大授粉的几率）,增加了受精的机会。相对而言,具有高效访花者的两个居群,资源较     

华北蓝盆花的开花特性及传粉生态学研究

华北蓝盆花具头状花序 ,花冠蓝紫色 ,1朵花的花期约为 7～ 8d。边缘花比中部花大 ,但二者在花药大小、花丝长度、花柱长度、柱头直径等方面无显著差异。花粉寿命约为 4 d,花柱在花开后逐渐伸长 ,柱头微二裂、干燥型。同一花序内的花朵在 2～ 3d内逐渐开放 ,晚开的花散粉时、早开的花花柱仍未伸长 ,因此 ,花药与柱头之间有空间间隔和时间间隔 ,同一花序内的各花朵间相互传粉的机会很低。按照杂交指数 ,其繁育系统属于异交、部分自交亲和、需要传粉者。依据 P/ O数据 ,其繁育系统属于兼性异交。罩网试验显示 ,华北蓝盆花没有自发的自花传粉现象 ,自然条件下也没有无融合生殖现象 ,没有访花者访问便不能座果。在华北蓝盆花花序上记录到的访花者有 10余种 ,以双翅目、鞘翅目、膜翅目昆虫为主 ,最常见的访花者是雏蜂虻和豆芫菁。开花后不同日期 ,访花者的访花频率有所不同 :开花第 2天到第 7天的花序上 ,访花者的访花频率较高 ;开花第 4天 ,花序上访花者的访花频率最高。同一天内的不同时间 ,访花者的访花频率也有差异 ,一般规律是 :11:0 0～ 15 :0 0之间 ,访花者的访花频率较高。此外 ,气温骤降、阴天及降雨能使访花者的访花频率明显降低。     

Nectary Structure, Nectar Secretion Patterns and Nectar Composition in Two Helleborus Species

Abstract: The morphological and cytological characteristics of nectaries of Helleborus foetidus and H. bocconei during the secretory period are reported. The nectaries are derived from modified petals and secrete nectar continuously for about 20 days; they consist of a single layered epidermis, nectar-producing parenchyma and photosynthesizing parenchyma. Nectar secretion is holocrine and the nectar is released by rupture of the wall and cuticle of each epidermal cell. The nectaries of the two species differ in number and external morphology. In H. foetidus, secretion begins before anthesis and secretion rate decreases with nectary age. In H. bocconei it begins on the day of anthesis and proceeds at a constant rate. The nectar has a high sugar content, mainly sucrose, and also contains lipids and proteins.     

长白山针阔混交林紫椴的泌蜜量

紫椴是东北东部地区原始针阔混交林中优势种或共优种之一,是优质的用材树种,更是重要的蜜源植物。本文研究长白山地区紫椴从单株到种群(林分)的泌蜜量,建立单株泌蜜量回归模型,并估计林分尺度泌蜜量,分析紫椴泌蜜量与胸高断面积以及材积或蓄积之间的关系。结果表明: 紫椴单花开花时间为6～8 d,泌蜜时间为5 d,平均每朵花总泌蜜量为8.58 mg。花蜜的糖浓度在一天中有变化,中午高于早晨和傍晚,平均糖浓度为37.7%。样地紫椴的平均胸径为40 cm,单株开花量为18万个,泌蜜量为1.56 kg(或纯糖0.588 kg)。每公顷紫椴的泌蜜量为79～147 kg(或0.0686～0.1285 m³,纯糖29.78～55.42 kg)。林分泌蜜量与总断面积以及蓄积量密切相关,可以利用森林资源调查数据估算紫椴泌蜜量。     

Removal Effects on Nectar Production in Bat‐pollinated Flowers of the Brazilian Cerrado

We tested the removal effect on nectar production in four bat‐pollinated cerrado flowers. We compared the amount of nectar after 7–12 removals with the accumulated nectar in non‐manipulated flowers after 12 h. In all, but one species, removals increased volume by 1.5–4.6 times and sugar content by 1.6–4 times, which may affect flower visitation by bats, pollen flow, and reproduction. Abstract in Portuguese is available in the online version of this article.     

Nectar biodiversity: a short review

 Nectaries differ in many aspects but a common feature is some kind of advantage for the plant conferred by foraging of consumers which may defend the plant from predators in the case of extrafloral nectaries, or be agents of pollination in the case of floral nectaries. This minireview is concerned mainly with floral nectaries and examines the following characteristics: position in flower; nectary structure; origin of carbohydrates, aminoacids and proteins; manner of exposure of nectar; site of nectar presentation; volume and production of nectar in time; sexual expression of flower and nectary morphology; nectar composition and floral sexual expression; variability of nectar composition; fate of nectar; energy cost of nectar production. The species of certain large families, such as Brassicaceae, Lamiaceae and Asteraceae, resemble each other in nectary organisation; other families, such as Cucurbitaceae and Ranunculaceae, have various types of organisation. A scheme is presented to illustrate factors influencing nectary and nectar biodiversity. Received July 23, 2002; accepted September 18, 2002 Published online: June 2, 2003     

Nectar Dynamics and Sugar Composition in Flowers of Silene and Saponaria Species (Caryophyllaceae)

Abstract: Nectar production in Saponaria officincilis and in five species of Silene (S. ciba, S. dioica, S. noctiflora, S. nutans, S. vulgaris ) was examined during two consecutive years (May to July 1993, and May to June 1994) in the Botanical Garden of the University of Giessen. Nectar volume and sugar concentration were studied in relation to time of day, flower sex, flower age, and flowering stage. Nectar amount in all species studied (except S. dioica ) increased in the afternoon or in the evening until midnight (or until the early morning in S. nutans ). After midnight and until midday, nectar volume in non-visited flowers (except S. dioica ) decreased. Nectar volume in non-visited S. dioica flowers increased constantly with flower age, indicating a stable nectar secretion rate, possibly favouring both day- and night-active flower visitors. Even at the time of highest nectar secretion, all species studied presented several nectarless flowers. Sucrose dominance in the nectar of the nocturnal species S. nutans and Saponaria officinalis fits well with the general syndrome of flowers pollinated by hawkmoths. The syndrome also applies to the nocturnal but regularly selfing, S. noctiflora . The more generalis-tic species S. dioica and S. vulgaris , which are regularly visited by bumblebees as well as nocturnal moths, secreted hexose-domi-nant nectar. Unexpectedly, Silene alba , the only nocturnal species that strictly excluded day-active flower visitors by closing flowers during the day, also secreted hexose-dominant nectar. In some cases, nectar volumes and nectar concentration differed significantly between hermaphroditic, male, and female flowers. Female flowers of S. alba, S. dioica , and S. nutans contained significantly less concentrated nectar than male or hermaphroditic ( S. nutans ) ones. In S. noctifiora and S. vulgaris the difference was not statistically significant but nectar concentration did show the same tendency.     

Nectar sugar composition and volumes of 47 species of Gentianales from a southern Ecuadorian montane forest

BACKGROUND AND AIMS: This study investigates 47 taxonomically related species (Gentianales), all native to a tropical montane forest in southern Ecuador, in terms of nectar chemistry and nectar volumes in relation to pollination biology. METHODS: Nectar volumes of covered (24-h production) and uncovered (standing crop) flowers were measured in the natural habitat. Sucrose, fructose and glucose were quantified in the nectar using high performance liquid chromatography. Flower visitors were observed. KEY RESULTS: Nectar sugar concentration did not differ significantly among the pollination syndromes. Regarding sugar composition, the only significant differences were found in chiropterophilous and myiophilous flowers, which had a significantly lower sugar ratio than sphingophilous flowers. A separation of chiropterophilous and myiophilous flowers from the other pollination syndromes is further substantiated by non-linear multidimensional scaling using the chord-normalized expected species shared index of dissimilarity based on nectar sugar compositions. The matrix test revealed no correlation of observed floral visitors to nectar concentrations; however, a weak significant correlation was found between floral visitors and nectar sugar compositions. The nectar volumes of covered and uncovered flowers are related to, and differ significantly among, pollination syndromes. Matrix tests revealed correlation between floral visitors and nectar volume of covered flowers and, to a lesser extent, of uncovered flowers. CONCLUSIONS: Sucrose is the predominant floral nectar sugar in the order Gentianales, suggesting that nectar sugar composition is a conservative characteristic. However, some degree of an adaptive convergence of floral nectar compositions to principal pollinator type within the constraints set by phylogenetic history is likely. The driving force to visitation appears to be the volume of nectar the visitor can expect to consume.     

Nectar yeasts in Delphinium nuttallianum (Ranunculaceae) and their effects on nectar quality

Microorganisms colonize the nectar of many angiosperms. Variable diversity and spatio-temporal dynamics of nectar-inhabiting microorganisms (e.g., yeasts) may drive variation in nectar sugar composition and subsequent plant–pollinator interactions. We assessed yeast frequency of occurrence and density in the nectar of the perennial herb, Delphinium nuttallianum, across multiple spatio-temporal scales, including flower lifetime and sex-phase transition, flowering season, populations, and years. We tested the hypothesis that pollinators vector yeasts by comparing densities between virgin flowers and those open to visitation. Finally, we identified yeasts using molecular methods and tested for an association between yeast density and nectar composition using ultra-performance liquid chromatography. Yeasts were frequent colonists of Delphinium nectar, occurring in all populations and years sampled. Yeast frequency of occurrence and density varied across most spatio-temporal scales examined. Pollinators were vectors of yeast: virgin flowers remained yeast-free, while those open to visitation became inoculated. Nectar samples were species-poor, with a majority colonized by Metschnikowia reukaufii. Finally, increasing yeast density was correlated with a decrease in sucrose and an increase in monosaccharides. Our results document that yeasts form species-poor communities in populations of this hermaphroditic perennial, in addition to highlighting their spatio-temporal dynamics and effects on nectar quality. Spatio-temporal variation in frequency of occurrence, density, and changes in nectar may have important implications for the nature and strength of interactions between Delphinium and its pollinators.     

Postpollination Nectar Reabsorption and Its Implications for Fruit Quality in an Epiphytic Orchid1

We present evidence that pollination triggers nectar reabsorption in flowers of the epiphytic orchid Mystacidium venosum. The amount of sugar in nectar of M. venosum decreased significantly by more than 50 percent within 72 hours of pollination. Hand–pollinated flowers from which nectar was previously removed set significantly smaller fruits with a lower percentage of viable seeds than hand–pollinated flowers containing nectar, suggesting that resources reclaimed by nectar resorption are allocated to fruit production.     

Nectar Production of Two Sympatric Species of Columnar Cacti

We examined the potential for exploitation competition by differential attractiveness in two sympatric species of columnar cacti on Curaçao, Netherlands Antilles. Stenocereus griseus and Subpiloccreus repandus have temporally overlapping flowering phenologies and share pollinators. We examined nectar volume, energy contents, and sugar ratios for the entire night and at two-hour intervals in both cactus species. Except for a burst in nectar volume and sugar concentration by Subpilocereus repandus during the first two hours of anthesis, nectar secretion patterns, energy contents, and sugar ratios (70% hexose) were similar for the two species. The standing crops of nectar in both species were kept very low by bat pollinators. We suggest that the potential for exploitation competition between Stenocereus griscus and Subpilocereus repandus is currently very low on Curapo.     

Nectar Secretion Pattern and Removal Effects in Six Argentinean Pitcairnioideae (Bromeliaceae)

Nectar secretion pattern and effects of nectar removal were analyzed in six hummingbird-pollinated Argentinean Pitcairnioideae: Abromeitiella brevifolia. A. lorentziana, Deuterocohnia longipetala. Dyckia floribunda, D. ragonesei and Puya spathacea. Flower lifetime was determined in each case. Nectar volume, concentration and sugar production were measured from bud opening till fading of flowers at different time intervals according to the species' flower life-span. Nectar volume varies while nectar concentration and cumulative sugar production increase as a function of flower age in all the studied species. In general, total nectar volume and concentration values can be affected by periodic removal, but total sugar production is unaffected. Only in Puya spathacea did periodic nectar harvesting reduce the total amount of sugar produced.