Floral nectary structure and nectar composition in Eccremocarpus scaber (Bignoniaceae), a hummingbird-pollinated plant of central Chile

Surface features, anatomy, and ultrastructure of the floral nectary of Eccremocarpus scaber (Bignoniaceae), pollinated predominantly by the largest-known hummingbird (Patagona gigas gigas), were studied together with nectar sugar content and secretion rate. The annular disk nectary comprises epidermis, secretory and ground parenchyma with intercellular spaces, and branched vascular bundles terminating in the secretory parenchyma where only phloem is found. Amyloplasts and vacuoles increase in size throughout development, the latter becoming sites of organelle degradation. Transferlike cells in nectary phloem and P-proteinlike fibrillar material in phloem parenchyma were observed. Flowers produced around 32 μl of nectar (mostly after anthesis) with 11 mg of sugar composed of fructose, glucose, sucrose, and maltose in a ratio of 0.34:0.32:0.17:0.17. Morphological studies as well as the presence of maltose and glucose in nectar suggest storage of the originally phloem-derived sugars as starch with its subsequent hydrolysis. The low sucrose/hexose ratio (0.25) and high nectary secretion force (nectar per flower biomass) observed places E. scaber close to large-bodied bat-pollinated plants. A hypothesis based on nectar origin and nectar secretion is advanced to explain pollinator-correlated variation in sucrose/hexose ratio.     

Four o'clock pollination biology: nectaries,nectar and flower visitors in Nyctaginaceae from southern South America

Floral nectary structure and nectar sugar composition were investigated in relation to other floral traits and flower visitors in contrasting species of Nyctaginaceae from southern South America, representing four tribes (Bougainvilleeae, Colignonieae, Nyctagineae, Pisoneae). Our comparative data will aid in the understanding of plant–pollinator interactions and in the development of hypotheses on the origin of floral and reproductive characters in this family. The nectaries are located on the inner side of the staminal tube. The nectariferous tissue is composed of an epidermis and three to ten layers of secretory parenchymal cells, supplied indirectly by the filament vascular bundles. Stomata appear to be associated with nectar secretion. For the first time in Nyctaginaceae, nectary ultrastructure is described in Boerhavia diffusa var. leiocarpa. Nectary parenchyma cells are densely cytoplasmic and contain numerous starch grains. Plasmodesmata connect the nectariferous cells. Flowers of Nyctaginaceae secrete a small volume of nectar of variable concentration (10–47%). Nectar is dominated by hexoses, but Mirabilis jalapa showed a balanced proportion of sucrose and hexoses. Hymenoptera are the most common visitors for most species; nocturnal Lepidoptera are the most common visitors for M. jalapa and Bougainvillea stipitata. We found relatively low variation in the nectary characteristics of Nyctaginaceae compared with broad variation in flower structure, shape, colour and nectar traits. © 2013 The Linnean Society of London     

短果大蒜芥花蜜腺的发育解剖学研究

短果大蒜芥（Sisymbrium loeselii L.var.brevicarpum Z.X.An)花蜜腺位于雄蕊基部花托上,属十字花科环状花蜜腺类型中的侧棱环四圆环亚型。蜜腺由分泌表皮,产蜜组织和维管束组成。分泌表皮上有变态气孔器,蜜腺中部的气孔器呈舟状分布。产蜜组织中的维管束来自于花托中的维管束分支,属较进化的十字花科花蜜腺的亚型类型。蜜腺原基是在花的各部分原基分化后,由雄蕊基部花托表面区域的2－3层细胞,经反分化形成,环状蜜腺发生发育同步,在蜜腺的发育过程中,蜜腺组织中的液泡和淀粉粒都发生了有规律的变化,其原蜜汁由维管束提供,运转至产蜜组织,最后由变态气孔泌出。     

Extrafloral Nectaries in Acacia mangium

Our microscopy studies describe the anatomy of extrafloral nectaries on the abaxial side of the basal part of every leaf stalks of Acacia mangium. The lens-like nectary expands with the development of the leafstalk, peaks at the stage at which the leafstalk itself has reached its mature size. The nectary is composed of numerous small parenchyma cells and a nectar cavity in which the nectar is pooled. Those small parenchyma cells are divided into nectariferous tissue and epithelial cells, which line the lumen of the nectar cavity, and secretes the nectar into the same. Each nectary is surrounded by several vascular bundles, which probably afford the nectar. In addition to the microscopic observation, the chemical constituents of the nectar are analyzed by NMR, and it mainly consists of sugars with 60 % sucrose, 25 % glucose and 15 % fructose.     

地椒花蜜腺发育的解剖学观察

通过显微和亚显微观察对地椒花蜜腺的发育进行了研究。地椒花蜜腺位于子房基部的花盘上,属于盘状蜜腺,新鲜时呈绿色。蜜腺由分泌表皮和泌蜜组织组成,分泌表皮为一层细胞,表皮细胞角质膜较厚,表皮上分布着大量的气孔器,气孔器突出于表面;泌蜜组织细胞多层。花盘中央有维管束通向子房,在维管束和泌蜜组织之间有４￣５层大型的薄壁细胞。蜜腺由花盘的表皮及其内侧相邻的细胞发育而来,在发育过程中,其细胞中的液泡和贮藏的淀粉     

Nectary and gender‐biased nectar production in dichogamous Chamaenerion angustifolium (L.) Scop. (Onagraceae)

In dichogamous plants, nectar characteristics (i.e. nectar amount and its composition) can differ between sexual phases. In the present study, we investigated the structural organization of the floral nectary, nectar production and carbohydrate composition in the protandrous Chamaenerion angustifolium (L.) Scop. (Onagraceae). The receptacular nectary consisted of an epidermis with numerous nectarostomata, several layers of photosynthetic secretory parenchyma, and subsecretory parenchyma. Nectariferous tissue was not directly vascularized and starch grains were rarely observed in the secretory cells, occurring exclusively in the guard cells of modified stomata. The nectar was released via nectarostomata. The floral nectar was hexose rich (32.8/39.1/28.1% glucose/fructose/sucrose) and the total concentration was constant throughout the anthesis (47% on average). However, contrasting patterns in nectar amount and carbohydrate composition between the floral sexual phases were observed. On average, female‐phased flowers produced 1.4‐fold more nectar than male‐phased flowers, and although the nectar was sucrose rich during the male phase, it was hexose rich during the female phase, suggesting sucrose hydrolysis.     

THE FOLIAR AND FLORAL NECTARIES OF TURNERA ULMIFOLIA L.

The floral and foliar nectaries of Turnera ulmifolia are specialized and are representative of others found in the Turneraceae. The foliar and floral nectary systems must be treated independently. Foliar nectaries are organized into a definite structure (composed of a base, rim, secretory tissue, modified epidermis) and are supplied with vascular tissue composed of both xylem and phloem. Nectar from foliar nectaries contained equal concentrations of glucose, fructose, and sucrose. Floral nectaries are an integral part of the basal portion of each filament. The nectariferous tissue is not supplied with vascular tissue and secretion lasts only a few hours. Nectar from these staminal nectaries yielded a sucrose-dominant nectar containing also fructose, glucose, an unknown, and a trace amount of melezitose. Observations of flowering confirmed the reported short duration of the individual flowers.     

Anatomical and ultrastructural changes of the floral nectary ofPisum sativum L. during flower development

Summary The floral nectary ofPisum sativum L. is situated on the receptacle at the base of the gynoecium. The gland receives phloem alone which departed the vascular bundles supplying the staminal column. Throughout the nectary, only the companion cells of the phloem exhibited wall ingrowths typical of transfer cells. Modified stomata on the nectary surface served as exits for nectar, but stomatal pores developed well before the commencement of secretion. Furthermore, stomatal pores on the nectary usually closed by occlusion, not by guard-cell movements. Pore occlusion was detected most frequently in post-secretory and secretory glands, and less commonly in pre-secretory nectaries. A quantitative stereological study revealed few changes in nectary fine structure between buds, flowers secreting nectar, and post-secretory flowers. Dissolution of abundant starch grains in plastids of subepidermal secretory cells when secretion commenced suggests that starch is a precursor of nectar carbohydrate production. Throughout nectary development, mitochondria were consistently the most plentiful organelle in both epidermal and subepidermal cells, and in addition to the relative paucity of dictyosomes, endoplasmic reticulum, and their associated vesicles, the evidence suggests that floral nectar secretion inP. sativum is an energy-requiring (eccrine) process, rather that granulocrine.Abbreviations ER endoplasmic reticulum - GA glutaraldehyde - SEM scanning electron microscopy     

Nectaries and male-biased nectar production in protandrous flowers of a perennial umbellifer <Emphasis Type="Italic">Angelica sylvestris</Emphasis> L. (Apiaceae)

Nectar is the most common floral pollinator reward. In dichogamous species, floral nectar production rates can differ between sexual phases. We studied the structure of nectaries located on the stylopodium and nectar production in protandrous umbellifer Angelica sylvestris. Our study species produced nectar in both floral sexual phases. Nectar sugar concentration was low (on average 22 ± 11 %, mean ± SD) and the nectar hexose rich and composed of sucrose, glucose, fructose and a small amount of amino acids, including β-alanine, a non-protein amino acid. Although nectar composition and sugar concentration varied little between floral sexual phases, nectar production showed a threefold reduction during the stigma receptive period. This is in contrast to other studies of Apiaceae that have reported female-biased nectar production, but in the direction predicted by plant sexual selection theory, suggesting that in pollen-unlimited species, floral rewards mainly enhance male reproductive success. The structure of the nectary was similar at the two sexual stages investigated, and composed of a secretory epidermis and several layers of nectariferous and subsecretory parenchyma. The nectary cells were small, had large nuclei, numerous small vacuoles and dense, intensely staining cytoplasm with abundant endoplasmic reticulum, mitochondria and secretory vesicles. They contained abundant resin-like material that may potentially act as defence against microbes. Starch was rarely observed in the nectary cells, occurring predominantly at the female stage and mainly in guard and parenchyma cells in close proximity to stomata, and in subsecretory parenchyma. The main route of nectar release in A. sylvestris seems to be via modified stomata.     

异株百里香花蜜腺的发育解剖学研究

异株百里香（Thymus marschallianus Willd）花蜜腺分布于子房基部的花托上,结构蜜腺盘状,成熟时膨大,环绕在花托外。蜜腺组织由分泌表皮、产蜜组织和维管束三部分组成;组织化学染色显示淀粉粒的积累是在蜜腺细胞发育的最初和最后,因此将其归为非淀粉型蜜腺。在发育的过程中细胞液泡化动态明显,而淀粉粒和多糖均不具有明显的消长变化;蜜汁是由韧皮部运转到泌蜜组织中的,再由表皮细胞的角质层渗到细胞外。     

荔枝花蜜腺发育解剖学研究

荔枝花蜜腺呈盘状,位于子房和花萼之间的花托上。花盘蜜腺由表皮、产蜜组织、维管束组成。蜜腺的原始细胞由花托表面的2~3层细胞脱分化产生。成熟蜜腺产蜜组织细胞含有淀粉粒,为淀粉型蜜腺,表皮细胞内无淀粉粒。产蜜组织出现分化:PAS反应颜色深的细胞成网状分布,与表皮下方的1~2层细胞相连,构成蜜汁的运输通道;颜色浅的细胞分布于网眼处。蜜腺表皮上的角质层波状皱折,有泌蜜孔。表皮毛主要起保护作用,大部分蜜汁通过泌蜜孔排出。     

鹅掌柴花蜜腺的发育解剖学研究

对鹅掌柴(Scheffler octophylla Harms．)花蜜腺的发育进行解剖结构观察。鹅掌柴花盘蜜腺位于下位子房上方环绕花柱基部。蜜腺由分泌表皮、产蜜组织组成,心皮维管束与其相邻并发出一些伸入蜜腺基部的短分枝。蜜腺起源于心皮原基基部外侧的几层细胞。鹅掌柴花蜜腺为淀粉型蜜腺,淀粉粒为许多微小颗粒聚集成的复粒。原蜜汁由蜜腺基部维管束的筛管提供,达产蜜组织细胞和表皮细胞后以淀粉粒的形式贮藏。泌出的蜜汁一部分来自淀粉粒的降解,一部分来自泌蜜期输入的原蜜汁。表皮和产蜜组织细胞均具泌蜜功能。泌出的蜜汁大部分通过气孔排出,还有部分由角质层渗出。     

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     

A cell wall invertase controls nectar volume and sugar composition

Nectar volume and sugar composition are key determinants of the strength of plant–pollinator mutualisms. The main nectar sugars are sucrose, glucose and fructose, which can vary widely in ratio and concentration across species. Brassica spp. produce a hexose-dominant nectar (high in the monosaccharides glucose and fructose) with very low levels of the disaccharide sucrose. Cell wall invertases (CWINVs) catalyze the irreversible hydrolysis of sucrose into glucose and fructose in the apoplast. We found that BrCWINV4A is highly expressed in the nectaries of Brassica rapa. Moreover, a brcwinv4a null mutant: (i) has greatly reduced CWINV activity in the nectaries; (ii) produces a sucrose-rich nectar; but (iii) with significantly less volume. These results definitively demonstrate that CWINV activity is not only essential for the production of a hexose-rich nectar, but also support a hypothetical model of nectar secretion in which its hydrolase activity is required for maintaining a high intracellular-to-extracellular sucrose ratio that facilitates the continuous export of sucrose into the nectary apoplast. The extracellular hydrolysis of each sucrose into two hexoses by BrCWINV4A also likely creates the osmotic potential required for nectar droplet formation. These results cumulatively indicate that modulation of CWINV activity can at least partially account for naturally occurring differences in nectar volume and sugar composition. Finally, honeybees prefer nectars with some sucrose, but wild-type B. rapa flowers were much more heavily visited than flowers of brcwinv4a, suggesting that the potentially attractive sucrose-rich nectar of brcwinv4a could not compensate for its low volume.     

Floral nectar production and carbohydrate composition and the structure of receptacular nectaries in the invasive plant <Emphasis Type="Italic">Bunias orientalis</Emphasis> L. (Brassicaceae)

The data relating to the nectaries and nectar secretion in invasive Brassicacean taxa are scarce. In the present paper, the nectar production and nectar carbohydrate composition as well as the morphology, anatomy and ultrastructure of the floral nectaries in Bunias orientalis were investigated. Nectary glands were examined using light, fluorescence, scanning electron and transmission electron microscopy. The quantities of nectar produced by flowers and total sugar mass in nectar were relatively low. Total nectar carbohydrate production per 10 flowers averaged 0.3 mg. Nectar contained exclusively glucose (G) and fructose (F) with overall G/F ratio greater than 1. The flowers of B. orientalis have four nectaries placed at the base of the ovary. The nectarium is intermediate between two nectary types: the lateral and median nectary type (lateral and median glands stay separated) and the annular nectary type (both nectaries are united into one). Both pairs of glands represent photosynthetic type and consist of epidermis and glandular tissue. However, they differ in their shape, size, secretory activity, dimensions of epidermal and parenchyma cells, thickness of secretory parenchyma, phloem supply, presence of modified stomata and cuticle ornamentation. The cells of nectaries contain dense cytoplasm, plastids with starch grains and numerous mitochondria. Companion cells of phloem lack cell wall ingrowths. The ultrastructure of secretory cells indicates an eccrine mechanism of secretion. Nectar is exuded throughout modified stomata.     

Developmental and Anatomical Studies on the Floral Nectaries of Capsella bursa-pastoris

The development and structure of the floral nectaries of Capsella bursa-pastoris (L.) Medic. were examined. The nectaries consisted of four separated parts which were semiorbicular and were morphologically and anatomically similar to one another. They were located at the receptacle between stamens, and each part was composed of secretory epidermis, nectariferous tissue and vascular bundles, belonging to structural nectary. When the various floral organs were developed, 2--3 superficial layer cells of the receptacle between stamens became meristemoid and contributed to primordia the formation of nectary. By intercalary meristematic activity, the four nectaries formed synchronously. During the different stages of nectary differentiation, the content of starch gra ins and vacuolation in the cells of epidermis and nectariferous tissue changed regularly. According to the structural and histochemical changes the pre-nectar might be supplied by phloem. The nectar formed in nectariferous tissue was then secreted to the sub-stomatal chamber and where it was finally excreted from the stoma.     

荆条花蜜腺发育解剖学研究

荆条（Ｖｉｔｅｘ ｃｈｉｎｅｎｓｉｓ Ｍｉｌｌ．）花蜜腺属于淀粉型子房蜜腺,呈圆筒状环绕于子房的基部。蜜腺外观上无特殊结构,表面有。由分泌表皮和泌蜜组织组成,包括分泌表皮、气孔器、泌蜜薄壁组织和维管束。密腺和子房壁起源相同。花蕾膨大期,泌蜜组织细胞中产生大液泡;露冠期,泌蜜组织中形成维管束;花蕾初放期,分泌表皮细胞分化形成气孔器,无气孔下室,淀粉粒的积累在此期达到高峰;盛花期,蜜腺中已无淀粉粒,密     

Structure of the receptacular nectary and circadian metabolism of starch in the ant‐guarded plant Ipomoea cairica (Convolvulaceae)

Nectaries occur widely in Convolvulaceae. These structures remain little studied despite their possible importance in plant–animal interactions. In this paper, we sought to describe the structure and ultrastructure of the receptacular nectaries (RNs) of Ipomoea cairica, together with the dynamics of nectar secretion. Samples of floral buds, flowers at anthesis and immature fruits were collected, fixed and processed using routine methods for light, scanning and transmission electron microscopy. Circadian starch dynamics were determined through starch measurements on nectary sections. The secretion samples were subjected to thin layer chromatography. RNs of I. cairica were cryptic, having patches of nectar‐secreting trichomes, subglandular parenchyma cells and thick‐walled cells delimiting the nectary aperture. The glandular trichomes were peltate type and had typical ultrastructural features related to nectar secretion. The nectar is composed of sucrose, fructose and glucose. Nectar secretion was observed in young floral buds and continued as the flower developed, lasting until the fruit matured. The starch content of the subglandular tissue showed circadian variation, increasing during the day and decreasing at night. The plastids were distinct in different portions of the nectary. The continuous day–night secretory pattern of the RNs of I. cairica is associated with pre‐nectar source circadian changes in which the starch acts as a buffer, ensuring uninterrupted nectar secretion. This circadian variation may be present in other extrafloral nectaries and be responsible for full daytime secretion. We conclude that sampling time is relevant in ultrastructural studies of dynamic extranuptial nectaries that undergo various changes throughout the day.     

The systematic significance of floral morphology,nectaries, and nectar concentration in epiphytic cacti of tribes Hylocereeae and Rhipsalideae (Cactaceae)

A long-standing interest in cactus taxonomy has existed since the Linnaean generation, but an appreciation of the reproductive biology of cacti started early in the 1900s. Numerous studies indicate that plant reproductive traits provide valuable systematic information. Despite the extensive reproductive versatility and specializations in breeding systems coupled with the striking floral shapes, the reproductive biology of the Cactaceae has been investigated in approximately 10% of its species. Hence, the systematic value of architectural design and organization of internal floral parts has remained virtually unexplored in the family. This study represents the most extensive survey of flower and nectary morphology in the Cactaceae focusing on tribes Hylocereeae and Rhipsalideae (subfamily Cactoideae). Our objectives were (1) to conduct comparative morphological analyses of flowers and floral nectaries and (2) to compare nectar solute concentration in these two tribes consisting of holo- and semi-epiphytic species. Flower morphology, nectary types, and sugar concentration of nectar have strong taxonomic implications at the tribal, generic and specific levels. Foremost, three types of nectaries were found, namely chamber nectary (with the open and diffuse subtypes), furrow nectary (including the holder nectary subtype), and annular nectary. All Hylocereeae species possess chamber nectaries, in which the nectarial tissue has both trichomes and stomata. The Rhipsalideae are distinguished by two kinds of floral nectaries: furrow and annular, both nectary types with stomata only. The annular nectary type characterizes the genus Rhipsalis. Nectar concentration is another significant taxonomic indicator separating the Hylocereeae and Rhipsalideae and establishing trends linked to nectar sugar concentration and amount of nectar production in relation to flower size. There is an inverse relationship between flower size and amount of nectar production in the smaller Rhipsalideae flowers, in which nectar concentration is more than two-fold higher despite the smaller volume of nectar produced when compared to the large Hylocereeae flowers. Variability of nectary morphology and nectar concentration was also evaluated as potential synapomorphic characters in recent phylogenies of these tribes. In conclusion, our data provide strong evidence of the systematic value of floral nectaries and nectar sugar concentration in the Cactaceae, particularly at different taxonomic levels in the Hylocereeae and Rhipsalideae.     

芝麻菜花蜜腺的结构和发育研究

通过解剖镜观察、石蜡切片和薄切片等方法,对芝麻菜的花蜜腺的位置、形态、结构、发育过程及泌蜜前后组织化学变化进行了研究。芝麻菜花蜜腺4枚,分成两对,其中一对侧蜜腺较大,棱柱状,分别着生在外轮2个短雄蕊基部内侧的花托上,结构上由表皮、产蜜组织和维管组织构成;另一对中蜜腺较小,近棒状,分别着生在内轮4个长雄蕊外侧的花托上,结构上仅由表皮和产蜜组织构成。二者表皮细胞外都具角质层,且蜜腺产蜜组织细胞中只含少量的多糖物质。两类蜜腺的蜜汁均由变态气孔泌出体外。无论侧蜜腺还是中蜜腺,蜜腺原基皆是在雌、雄蕊已分化后,由花托相应位置表皮下的1～2层细胞分裂形成的。在蜜腺发育中,产蜜组织细胞在泌蜜前后不具明显的液泡变化。