The Floral Nectary ofDigitalis purpureaL., Structure and Nectar Secretion

The floral nectary of the foxglove (Digitalis purpureaL.), locatedat the base of the ovary, was examined by: scanning electronmicroscopy; quantitative bright-field microscopy via computer-aided3-D reconstruction from serial sections; morphometric procedures;transmission electron microscopy and measurement of nectar effluxunder different experimental conditions. Time-lapse video recordingvia a microscope with incident light clearly showed that thenectar escaped from the apertures of modified stomata. The volumeflux via individual stomatal apertures was 0.31±0.1 nlmin^-1; therefore only a fraction of the total number of stomataper nectary (115±8) would be sufficient to dischargethe amount of nectar reported in previous publications. Thestomatal apertures are continuous with intercellular spacestraversing the small-celled nectariferous tissue. The latteris vascularized only by phloem, whose termini consists of rowsof slender cells. These sieve-like cells are surrounded by moreor less isodiametrical sheath cells with dimensions similarto the secretory cells. Details of nectary functioning are basedon enhanced structural information, complementary data on nectardischarge after experimental manipulations and the nature ofthe effluence.Copyright 1998 Annals of Botany Company Digitalis purpureaL.; foxglove; floral nectary; (ultra-)structure; 3-D reconstruction; morphometry; nectar flow; time-lapse video recording.     

多叶斑叶兰花蜜特征和分泌规律及其对昆虫访花的影响

为探讨多叶斑叶兰(Goodyera foliosa)花蜜体积和成分的变异性,及其对传粉昆虫的影响,采用毛细管法、折光仪和高效液相-蒸发光法对一天内不同时间段单花花蜜体积、总糖浓度和可溶性糖成分及含量进行检测,用摄像机对其传粉昆虫访花行为进行观察。结果表明,单花花蜜体积在10:00达到最大[(7.19±2.29)μL, n=10],糖浓度在16:00达到最大[(25.85±1.83)%,n=10]。花蜜中的主要可溶性糖为果糖和蔗糖,果糖在上午10:00含量达到最高(78.310 mg/g);蔗糖在下午16:00含量达到最高(247.600 mg/g)。中华蜜蜂(Apis cerana)访花时间为每天9:00-15:00,访花高峰期在10:00-14:00;橘尾熊蜂(Bombus friseanus)访花时间为每天8:00-16:00,访花高峰期在10:00-12:00和16:00-18:00。因此,这两种传粉昆虫访花频率与多叶斑叶兰单花花蜜体积及糖浓度的变化有一定的相关性,它们更倾向于访问较高花蜜体积及糖浓度的花朵。多叶斑叶兰花蜜成分中蔗糖占优势,能有效吸引各种蜂类、蝶类和蚁类访花,且访问频率较高,中华蜜蜂和橘尾熊蜂能携带其花粉块,是主要的有效传粉昆虫。花形态和访花昆虫的体型大小的匹配,决定了是否成为有效传粉昆虫。     

Floral Nectaries of Rosmarinus officinalis L. Structure, Ultrastructure and Nectar Secretion

The nectary of Rosmarinus officinalis L. has the form of a four-lobed,asymmetrical disc situated around the base of the ovary. Thenectary lobe facing the lower flower lip is enlarged and isthe only one to have modified stomata. Vascular strands consistingof phloem only occur in the nectariferous tissue. It is suggestedthat the pre-nectar originating in the phloem accumulates primarilyas starch grains in plastids of the nectariferous cells. Thenumber of grains is very large before anthesis and decreasesconsiderably at anthesis. The transport of the pre-nectar tothe various nectariferous cells appears to be mainly via thesymplast. It could not be determined whether the process ofelimination of the nectar is solely eccrine or partly granulocrine. Rosmarinus officinalis, nectary, nectar secretion, starch grains, phloem     

The Floral Nectary of Tropaeolum majus L.--The Nature of the Secretory Cells and the Manner of Nectar Secretion

The floral nectary of Tropaeolun majus L. was studied with theaid of a microscope with transmitting and incident light, atransmission electron microscope and a scanning electron microscope.The Gomori method was used for the localization of acid phosphatase.As a result of this investigation the previously accepted viewthat nectar in this plant is secreted only from the hair tipsof the inner epidermis of the calyx spur was found to be inaccurate.The present studies showed that the parenchyma cells locatedbetween the inner epidermis and the region of the vascular bundlesof the lowest third of the spur, are the main nectar-secretingelements of the nectary. These secretory cells release the nectarsolution into intercellular spaces leading to modified stomata,through which it is exuded into the spur cavity. The modifiedstomata occur in the lowest portion of the spur only. At thestage of secretion small droplets of liquid of high viscositywere observed on the epidermal hairs. These droplets presumablycontain polysaccharides and a certain amount of sugar.     

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.     

Intraspecific Variation in the Amino Acid Content of Floral Nectar

In order to evaluate the constancy of amino acid composition and concentration, several randomly collected samples of genuine floral nectar from different flowers of several plant species were analysed. Although there seems to exist a rough pattern of species-specific nectar amino acid composition, amino acid concentration varied strongly from sample to sample. Apart from the lack of constancy in concentration of floral amino acids, our samples do not show amino acid concentration values as assigned typical for the pertinent pollination mode by other workers.     

The Importance of Osmosis in Nectar Secretion and its Consumption by Insects

SYNOPSIS. Nectar is a simple food consisting of varying proportionsof sucrose, glucose and fructose dissolved in water. The mechanismsof its secretion are poorly understood. Osmosis may explainwhy hexose-rich nectars are produced in larger volumes and aremore dilute than sucrose-rich nectars. Unless protected, nectartends to equilibrate with ambient humidity, and the concentrationsavailable to flower visitors can vary from 7–70% w/w.Most nectars are osmotically concentrated, especially when richin hexose sugars. The only digestion needed is sucrose hydrolysis,and monosaccharides and water are rapidly absorbed across themidgut of insects. Large flying insects feeding on nectar producean excess of water which must be eliminated by evaporation andexcretion. Animals which utilise this attractively packagedand easily digested food source may have osmoregulatory problemswhen nectar concentrations do not match their water requirements.     

Floral Nectar, Nectary Structure and Pollinators in Some Argentinean Bromeliaceae

The floral nectar chemical composition and nectary structureof some Argentinean Bromeliaceae were studied, including fieldobservations on pollinators. Twenty species belonging to eightgenera from the three subfamilies were analysed. The nectarcomponents report is mostly new since no comprehensive studyhas been carried out on the family previously. Sugars were alwayspresent, while alkaloids, lipids, phenols, and proteins werenot detected in any sample. Reducing acids were found in threespecies. Amino acids were detected in a very low concentrationin only about half the samples. Pitcairnioideae species showa mean balanced disaccharide/monosaccharide nectar sugar composition,Bromelioideae had hexose-rich nectars and Tillandsioideae saccharose-dominantones. Nectar concentration ranged from 16 to 48%. All taxa bearseptal nectaries with many common features. Pitcairnioideaeand Tillandsioideae members have half-inferior ovaries, a featuremostly overlooked in previous studies. Three types of nectaryarchitecture were recognized in both subfamilies. Bromelioideaehave inferior ovaries and possess comparable nectaries. Hummingbirdsconstitute the main flower pollinators of many species but butterfliesand bees were occasionally seen in two species, cropping nectarand pollen, respectively. Argentinean Bromeliaceae, floral nectar, nectary structure, pollinators, Abromeitiella, Aechmea, Bromelia, Deuterocohnia, Dvckia, Puva, Tillandsia, Vriesea     

Floral Nectar, Nectary Structure and Pollinators in Some Argentinean Bromeliaceae

The floral nectar chemical composition and nectary structureof some Argentinean Bromeliaceae were studied, including fieldobservations on pollinators. Twenty species belonging to eightgenera from the three subfamilies were analysed. The nectarcomponents report is mostly new since no comprehensive studyhas been carried out on the family previously. Sugars were alwayspresent, while alkaloids, lipids, phenols, and proteins werenot detected in any sample. Reducing acids were found in threespecies. Amino acids were detected in a very low concentrationin only about half the samples. Pitcairnioideae species showa mean balanced disaccharide/monosaccharide nectar sugar composition,Bromelioideae had hexose-rich nectars and Tillandsioideae saccharose-dominantones. Nectar concentration ranged from 16 to 48 %. All taxabear septal nectaries with many common features. Pitcairnioideaeand Tillandsioideae members have half-inferior ovaries, a featuremostly overlooked in previous studies. Three types of nectaryarchitecture were recognized in both subfamilies. Bromelioideaehave inferior ovaries and possess comparable nectaries. Hummingbirdsconstitute the main flower pollinators of many species but butterfliesand bees were occasionally seen in two species, cropping nectarand pollen, respectively. Argentinean Bromeliaceae,, floral nectar, nectary structure, pollinators, alkalinity, abromeitiella, Aechmea, Bromelia, Deuterocohnia, Dyckia, puya, Tillandsia, vriesea     

Species Richness of Yeast Communities in Floral Nectar of Southern Spanish Plants

Floral nectar of insect-pollinated plants often contains dense yeast populations, yet little quantitative information exists on patterns and magnitude of species richness of nectar-dwelling yeasts in natural plant communities. This study evaluates yeast species richness at both the plant community and plant species levels in a montane forest area in southern Spain, and also explores possible correlations between the incidence of different yeast species in nectar and their reported tolerance to high sugar concentrations, and between yeast diversity and pollinator composition. Yeast species occurring in a total of 128 field-collected nectar samples from 24 plant species were identified by sequencing the D1/D2 domain of the large subunit rDNA, and rarefaction-based analyses were used to estimate yeast species richness at the plant community and plant species levels, using nectar drops as elemental sampling units. Individual nectar samples were generally characterized by very low species richness (1.2 yeast species/sample, on average), with the ascomycetous Metschnikowia reukaufii and Metschnikowia gruessii accounting altogether for 84.7% of the 216 isolates identified. Other yeasts recorded included species in the genera Aureobasidium, Rhodotorula, Cryptococcus, Sporobolomyces, and Lecythophora. The shapes and slopes of observed richness accumulation curves were quite similar for the nectar drop and plant species approaches, but the two approaches yielded different expected richness estimates. Expected richness was higher for plant species-based than for nectar drop-based analyses, showing that the coverage of nectar yeast species occurring in the region would be improved by sampling additional host plant species. A significant correlation was found between incidence of yeast species in nectar and their reported ability to grow in a medium containing 50% glucose. Neither diversity nor incidence of yeasts was correlated with pollinator composition across plant species.     

可变剪切在植物成花转换中的作用

精确调控成花转换,确保植物在适宜环境下开花,对于植物的成功繁殖和物种繁衍至关重要。开花由多种分子机制在转录、转录后和蛋白质水平进行调控。可变剪切(AS)是一种普遍的转录后水平调控过程,可从单个基因产生多个转录本,从而丰富转录组和蛋白质组的多样性。大量研究表明,可变剪切在成花转换过程中发挥重要作用。根据发育和环境条件, AS能够影响mRNA的稳定性和/或蛋白亚型的功能,从而调控开花相关基因的功能转录本和/或功能蛋白水平。揭示成花相关pre-mRNA的AS作用将进一步增进人们对开花相关基因功能以及整个成花转换调控网络的认识。该文归纳了涉及成花转换的AS研究进展,并针对各个调控途径进行总结,以期为进一步研究植物AS和成花转换调控机制提供参考。     

刺五加、短梗五加的花蜜分泌节律、花蜜成分及访花者多样性的比较研究

野外定位观测刺五加（Eleutherococcus senticosus）、短梗五加（E.sessiliflorus）的花蜜分泌节律、访花者的多样性,室内分析其花蜜的主要成分。结果表明,刺五加雄株的花杂在开花1－3（4)d分泌花蜜,雌株在开花5－7,6－8或7－9d分泌花蜜;短梗五加以及刺五加两性株的部分花杂,在开花后有两次分泌花蜜的过程：第1次与花药开裂散粉时间一致,第2次与柱头具可授性的时间一致。而且,刺五加和短梗五加都由动物帮助传粉,花蜜分泌的时间与多数访花者的访花时间一致,在一天之中,散出花粉的花朵分泌花蜜的时间早于接受花粉的花杂,这种时间差异应该是植物控制该 花者流向并导致传粉成功的关键。短梗五加与刺五加之间以及刺五加不同性别的植株之间,花蜜的成分及相对含量各有特点,但都以果糖和葡萄糖为主。在刺五加、短梗五加花朵上记录到的访花昆明分别为50余种和40余种,多数隶属于膜翅目、鳞翅目、鞘翅目和半翅目。其中膜翅目的胡蜂、马蜂、熊蜂,双翅目的食蚜蝇、寄蝇等是刺五加、短梗五加的常见访花者。     

Floral Nectar Secretion and Ploidy in Brassica rapa and B. napus (Brassicaceae). II. Quantified Variability of Nectary Structure and Function in Rapid-cycling Lines

Haploid, diploid and tetraploid lines ofBrassica rapaL. (syn.campestris),and allotetraploidB. napusL., were examined to determine theinfluence of ploidy on floral features, particularly nectarymorphology and anatomy, and to relate nectary structure to nectarproduction capacity. Except for haploids, all lines were rapid-cycling.Average flower dry weight, and petal length and width, werein the descending orderB. napus>B. rapa (4n) >2n>n.Pollen grains of 4nplants were larger than those of 2nplants;haploids lacked pollen. All lines developed nectaries. Typically, each flower producedtwo pairs of nectaries, of different types and nectar productioncapacity. Normally, each lateral gland was located above thebase of a short stamen, and together this pair yielded mostof a flower 's nectar carbohydrate. Each median nectary aroseat the outer junction of the bases of two adjacent long stamens.All lateral nectaries received a vascular supply of phloem alone,but median glands received reduced amounts of phloem, or lackedvasculature altogether. Most nectaries were solitary, but 14%of all flowers, and especially those of 2n B. rapa,had at leastone median and lateral gland connected. Obvious variation existed in nectary morphology between ploidylevels, between flowers of the same plant, and even within flowers.Ten forms of each nectary type were recognized. Plants producingthe most nectar carbohydrate had high frequencies of lateralnectaries which were symmetrical, unfurrowed swellings. TetraploidsofB. rapahad both the highest frequencies of furrowed lateralglands, and of isolated segments of nectarial tissue at thatposition. Even these separated nectarial outgrowths receivedphloem and produced a nectar droplet. At the median location,nectaries were commonly of two forms: peg- or fan-shaped. Lobeson median nectaries, up to four per nectary, were detected inalmost half of glands of 4nflowers examined; lobes were absentin haploids. Brassica rapa; Brassica napus; flower size; nectar production; nectary variability; petal size; ploidyphloem; pollen; rapeseed     

Characteristics of Nectar Secretion by the Extrafloral Nectaries of Ricinus communis

The characteristics of nectar secretion by excised extrafloralnectaries of Ricinus have been examined. Secreted nectar wasfound to contain three sugars: sucrose, glucose and fructose,with glucose and fructose occurring in a 1: 1 ratio. All threesugars supported secretion when used in the culture medium andthe yield of nectar sugar was found to be concentration-dependent.Other sugar sources failed to support secretion. Experimentsusing ¹⁴C-sugars and ¹⁴CO₂ fed to intact plants allowed themovement of sugars through the nectary to be examined. Sucrosesynthesis occurs when excised glands are fed glucose and thisoccurs very early in the transport through the nectary. Themain sugar transported was sucrose, with little hydrolysis occurringuntil the final step of secretion. There was no evidence thatsucrose hydrolysis occurs either by invertase in the nectaror by a microbial flora. Inhibitors of respiration were foundto inhibit secretion as did anaerobiosis. Temperature also hada marked effect, with a temperature coefficient of 1.8. However,secretion of sucrose was not affected by anaerobic conditions,low temperatures or inhibitors of respiration as markedly asthat of glucose and fructose. Electron microscopy revealed the presence of a thickened andheavily stained wall at the inner border of the secretory epidermallayer. This wall contained numerous plasmodesmata at a frequencyof 14 per µm² and may represent an apoplastic barrier.Light microscope cytochemistry revealed that acid phosphataseis primarily located in the nectiferous tissue, while ATPaseis concentrated in the epidermis. The possibility that the nectarycontains two pathways for sucrose secretion, both apoplasticand symplastic, is discussed. Key words: Invertase, nectary, plasmodesmata, Ricinus communis, sucrose     

荔枝花蜜分泌规律及可溶性糖组分和含量的分析

为了解荔枝(Litchi chinensis)花蜜的分泌规律和主要组分,对‘糯米糍’、‘桂味’和‘怀枝’3个主栽品种雄花和雌花的花蜜分泌模式进行研究,并测定花蜜中可溶性糖的组分和含量。结果表明,采样期间果园阴天和晴天气温差异不明显,但阴天的相对空气湿度显著高于晴天。总体上阴天的荔枝花蜜分泌量高于晴天,雌花的花蜜分泌量高于雄花,‘桂味’和‘糯米糍’的花蜜分泌量均高于‘怀枝’。晴天花蜜中的可溶性固形物含量高于阴天的,且在雌花中表现尤为明显。‘怀枝’花蜜中的可溶性固形物含量最高,可达37.7%,‘桂味’其次,‘糯米糍’最少(17.7%)。利用高效液相色谱检测,荔枝花蜜中主要可溶性糖组分为葡萄糖、果糖和蔗糖,以葡萄糖含量最高。晴天时‘怀枝’雌花花蜜中可溶性糖含量达450.36μg m L–1,显著高于另外两个品种。这为荔枝栽培和花蜜生产提供科学依据。     

Nectar Sugar Production across Floral Phases in the Gynodioecious Protandrous Plant Geranium sylvaticum

Many zoophilous plants attract their pollinators by offering nectar as a reward. In gynodioecious plants (i.e. populations are composed of female and hermaphrodite individuals) nectar production has been repeatedly reported to be larger in hermaphrodite compared to female flowers even though nectar production across the different floral phases in dichogamous plants (i.e. plants with time separation of pollen dispersal and stigma receptivity) has rarely been examined. In this study, sugar production in nectar standing crop and secretion rate were investigated in Geranium sylvaticum, a gynodioecious plant species with protandry (i.e. with hermaphrodite flowers releasing their pollen before the stigma is receptive). We found that flowers from hermaphrodites produced more nectar than female flowers in terms of total nectar sugar content. In addition, differences in nectar production among floral phases were found in hermaphrodite flowers but not in female flowers. In hermaphrodite flowers, maximum sugar content coincided with pollen presentation and declined slightly towards the female phase, indicating nectar reabsorption, whereas in female flowers sugar content did not differ between the floral phases. These differences in floral reward are discussed in relation to visitation patterns by pollinators and seed production in this species.     

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.