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The structure of floral nectaries of ivy (Hedera helix) was investigated under light, scanning, and transmission electron microscopes. The nectar of ivy is located on top of the inferior ovary forming a distinct undulating disc between the base of petals and the style. The ivy nectary represents open and persistent nectaries. During consecutive days of anthesis, nectaries change their colour from green to brown. The secretory epidermis is covered with a thick, deeply striated cuticle, and nectar is released by nectarostomata. Epidermal cells exhibited plastids with plastoglobules and a few starch grains as well as vacuoles containing anthocyanins, the content of which increased during the successive days of anthesis and nectar secretion. Multi-layered glandular parenchyma and, underneath, subglandular tissue are located under the epidermis. The nectary was supplied by vascular bundles with phloem and xylem. Numerous chloroplasts were visible in the cytoplasm of the external layers of glandular parenchyma; they were either typical with small starch grains or untypical with circular arrangement of thylakoids. Amyloplasts containing storage starch grains and numerous small vacuoles were present in the cells of deeper layers of the nectar-bearing tissue. Druses, flocculent residue, myelin figures and spherical deposits of unknown origin were visible in the gland parenchyma vacuoles.
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Ecophysiological aspects of nectar reabsorption

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A number of approaches, both direct and indirect, have shown that nectar is reabsorbed by numerous plant species, irrespective of the age or sex of the flower. Furthermore, reabsorption occurs regardless of whether or not the flower has been pollinated. Reabsorption helps to maintain concentration of nectar and their viscosity and thereby encourages continued visits by pollinators. Conversely, the capacity to vary concentration of nectar sugars may confer evolutionary advantage by encouraging visits by more than one kind of pollinator and this is particularly important in regions where there is a paucity of pollinators. A further important role of nectar reabsorption is the maintenance of the energy equilibrium of the plant. A number of studies have shown that nectar production involves considerable energy expenditure requiring as much as 37% of the plant's daily production of energy by photosynthesis. The increased metabolic costs incurred by the plant during nectar production and secretion can reduce its growth and reproduction during the following season. Reabsorption of nectar that has not been collected by pollinators enables the plant to conserve at least some of the energy reserved for the secretion of nectar. Sugars reabsorbed from nectar can be re-used for the development of fruit and ovules - processes which demand large quantities of sugar. Despite convincing evidence for the reabsorption of nectar, few detailed studies have addressed the transport and incorporation of reabsorbed sugars. One of the questions that remain to be answered is 'What is the cellular basis for nectar reabsorption by the nectary?'.
Most orchids do not produce floral food-rewards. Instead, they attract pollinators by mimicry or deceit. When present, the most common floral food-reward is nectar. To date, nectary structure has been described for only two species of Maxillaria sensu lato, namely Maxillariella anceps and Ornithidium coccineum (formerly Maxillaria anceps and M. coccinea, respectively). Here, we describe that of a third species, Ornithidium sophronitis (formerly Maxillaria sophronitis). This species possesses floral characters concomitant with ornithophily. A ‘faucet and sink’ arrangement is present, with nectar secreted by a protuberance on the ventral surface of the column, collecting between column and tepal bases. The nectary of O. sophronitis shares many features with that of O. coccineum. It has a single-layered epidermis and 3- 5 layers of small, subepidermal, collenchymatous, secretory cells. Beneath these occur 2-3 layers of larger, subsecretory, parenchymatous cells supplied by phloem. Nectary cell vacuoles contain osmiophilic material and proteinaceous intravacuolar bodies. Moreover, distension of the nectary cuticle occurs as nectar accumulates between it and the secretory epidermis. Subsecretory cells, however, have thinner walls and contain flocculent, intravacuolar precipitates that may be related to the presence of flavonoids. Since the floral and nectary structure of O. sophronitis is very similar to that of closely related Ornithidium coccineum, it may have evolved in like manner in response to similar pollinator pressures.
Extrafloral nectaries on the abaxial surface of stipules were investigated in the Vicia angustifolia, Vicia sativa, Vicia sepium and Vicia grandiflora. In V. angustifolia nectaries were also located on the calyx surface. Nectaries were consisted of secretory hairs and 2-3 layers of subepidermal cells. Secretory hair was built af four cells of head, one stalk cell and basal cell. Head cells showed character of transfer cells because of walls ingrowths and dense cytoplasm with numerous mitochondria.
The presence of ruderal and mid-field vegetation promotes conservation of biodiversity and provides an additional source of food for insect pollinators. The white deadnettle is a common synanthropic plant visited frequently by various groups of insects, as it is a source of pollen and nectar. In 2012–2013, in the city of Lublin (Poland), signalling and food attractants in L. album flowers were analysed using light and scanning electron microscopy. The pipetting method was used for determination of nectar abundance in the flowers, and the content of sugars in the nectar was assessed with the use of an Abbe refractometer. It was found that the white dead-nettle flowers emitted a fragrance and were equipped with nectar guides, and the corolla and stamens had glandular trichomes and papillae secreting essential oils. On the stamens, there are also non-glandular trichomes, which play a role of pollen presenters. The nectary in the L. album flower has a shape of an irregular disc partly surrounding the base of the ovary. The content of sugars in the nectar was 43%. Sugar and honey yields per ha were calculated, and the values obtained were 153 kg and 191 kg, respectively.
To date, the structure of the cuniculus nectary has not been studied in detail. Furthermore, the secretory mechanism of such nectaries has not been investigated. The present paper describes, for the first time, the structural organization and ultrastructure of the cuniculus nectary in the moth-pollinated orchid Brassavola flagellaris Barb. Rodr. This tubular structure is situated between the perianth tube and ovary and, in its possession of thick, cellulose cell walls, resembles the nectary of ornithophilous taxa. The presence of large secretory vesicles that fuse with the plasmalemma indicate that granulocrine nectar secretion occurs in this species. The lumen of the cuniculus is lined with unicellular hairs. However, the cuticle overlying the whole epidermal surface lining the lumen (both glabrous and pubescent regions) was coated with nectar residues and became distended and cracked, indicating that this entire tissue is probably involved in nectar secretion.
The nectaries anatomical features of nine species of Pomoideae subfamily Aronia melanocarpa Ell., Cotoneaster horizontalis Decne, C. praecox Vilm.-Andr., C. lucida Schlecht., Crataegus monogyna Jacq., C. coccinea L., C. crus-galli L., Sorbus aucuparia L., Sorbus intermedia Pers. were compared. The flower longitudinal sections by means of vibratome and semithin slides by use of ultramicrotome were made. The thickness of nectary epidermis, glandular layer and the structure of subglandular layer as well as the run of vascular bundles were studied. It was stated that the height of epidermis cells and the thickness of their wall were approximate in the most investigated taxons. Stomata occurred in the nectaries of all species. They were located on a level with the epidermis cells in Aronia and Sorbus genera, in small hollows in Cotoneaster and considerably below the level of epidermis cells in Crataegus. The thickness of glandular layer was approximate for each genera. Sorbus intermedia and the species of Crataegus genus wer distinguishable by the thickest nectaries. The concentration of brachysclereids occurred in subglandular tissue of Crataegus crus-galli and of Sorbus both species. Vascular bundles, stocking the nectary, mostly consisted of phloem and xylem, but in Aronia, Cotoneaster praecox and C. lucida only phloem was noticed. The dependence between thickness of glandular layer, studied anatomical features and the volume of secreted nectar is discussed.
In the flowers of Allium there are found septal nectaries; in particular species, their outlet can be located in different parts of the ovary. The inflorescences of these plants are a rich source of nectar for insects. The location and structure of septal nectaries in the flowers of Allium giganteum Regel were investigated. Light and scanning electron microscopy was used. The septal nectaries were found to be located in the lower part of the ovary and in the gynophore on which the ovary is borne. Nectar is secreted into the nectary slits from which it flows through the ducts to three openings located in the upper part of the gynophore, from whence it gets outside in the vicinity of the expanded parts of the filaments. Sugar concentration in the nectar of A. giganteum averaged 54.5%, while sugar weight per flower was determined to be 0.36 mg. When converted into sugar weight per inflorescence, numbering more than 2,000 flowers, it was 771.7 mg.
Ecological traits of Erica carnea L. flowers and the morphology of floral nectaries were investigated using stereoscopic, light and scanning electron microscopy. The nectary in the flowers of Erica carnea is located in the basal part of the ovary. It represents the gynoecial nectary type. It has the form of a yellow, ribbed ring with eight outgrowths, pointed towards the base, which alternately adjoin the stamen filaments. The height of the nectary is 400 μm and its thickness 200 – 250 μm. The parenchyma of the nectary is composed of 6 – 8 layers. Nectar secretion occurs through anomocytic stomata with a diameter of 17 μm. Guard cells are only found on the outgrowths of the nectary and they are situated most frequently at the level of other epidermal cells. During nectar secretion, a small degree of pore opening was observed. In the flowers of Erica carnea, secondary nectar presentation was found, with the nectar accumulating at the base of the fused corolla.
The first apparent nectarivorous weevil, Smicronyx squalidus, is documented on Desmanthus illinoensis. Although found feeding from extra-floral nectaries (EFN’s) located between the petioles of D. illinoensis, it is believed that S. squalidus merely supplements its diet with nectar, due to an apparent lack of mouthpart modifications for a more efficient uptake of fluids. Photographs of the adult S. squalidus, as well as the EFN’s on D. illinoensis, are provided.
In the years 2007-2008, the flowering biology of Malus sargentii, an ornamental apple tree native to Japan, was studied in the conditions of Lublin (Poland). The daily rate of flower opening, flowering duration and flower visitation by insects were determined. The amount of nectar produced per flower and sugar content in the nectar were investigated. The size of nectaries and the micromorphology of their surface were examined using light and scanning electron microscopy. It was found that the greatest amount of flowers opened between 11.00 and 13.00. During this time, the largest number of insects was observed in the flowers. Bees (90%) were predominant among the insects, with a much smaller number of bumblebees (6%) and butterflies (4%). The flower life span was 5 days. Over this period, the flower produced, on the average, 0.71 mg of nectar with an average sugar content of 32%. The nectaries of Malus sargentii are orange-yellow coloured and they represent the hypanthial type. Due to the protrusion of the nectariferous tissue, they are classified as automorphic nectaries. The surface of the epidermal cells of the nectary was distinguished by distinct cuticle folds. A small number of stomata were located only in the basal part of the nectary. At the beginning of flowering, all stomata were closed, but secretion traces were observed near well-developed outer cuticular ledges.
The investigations were carried out using light and scanning electron microscopy. The flowers of Elaeagnus commutata grow in clusters of 1-4 in the leaf axils. They are actinomorphic, four-lobed, with a single perianth that is yellow from the adaxial side, while the abaxial side is silvery-white. Peltate hairs of different structure are found on both surfaces of the sepals. The conical epidermal cells of the lobes are covered with a thick striated cuticle. Cylindrical hairs were observed on the edges of the lobes. Peltate hairs also grew on the style. The dish-shaped nectary gland is located at the base of the style. Nectar is secreted through numerous, evenly distributed stomata located above or at the level of other epidermal cells. Different stages of stomatal development are evidence of the asynchronous functioning of the stomata. The nectary consists of small epidermal cells and 5-6 layers of secretory parenchyma. The deeper layers of the gland are composed of larger cells of subglandular parenchyma in which vascular bundles supplying the nectary run. Honey bees were the main pollinators of silverberry. Ten silverberry flowers produced an average of 12 g of nectar with a sugar concentration in the 29.5-34.5% range. The weight of pollen produced by 10 flowers was 3.33 mg.
The micromorphology of selected elements of Echium vulgare L. fl owers was investigated, with special attention to the structure of the nectaries and the stigma of the pistil as well as types of trichomes occurring on the surface of the calyx. The nectary had the shape of an uneven disc located around the lower region of the four-parted ovary of the pistil. The glandular cells formed a tier with a height of 330 μm and a radial width of 144 μm. Nectar was secreted onto the nectary surface through anomocytic stomata located at the level of other epidermal cells. Most of the stomata were open, with a different dimension of the pore. Their largest number was observed at the base of the nectary, and 462 stomata were noted on the whole surface of the nectary. The cuticle on the surface of the guard cells formed fi ne, circular striae. The subsidiary cells formed striated cuticular ornamentation, with the striae arranged radially in the direction of the stoma, whereas on the surface of other epidermal cells the striae formed an arrangement with different directions. The epidermis on the surface of the stigma formed regularly arranged papillae with a fan-shaped, expanded upper part which had corrugated outer walls, whereas the base of the cell formed a widened small column. The epidermis of the abaxial part of the calyx was covered by numerous non-glandular trichomes of different length which were made up of one or several cells. The glandular trichomes in the epidermis of the calyx grew with smaller density compared to the protective trichomes, and they were composed of a 1-2-celled stalk and a glandular head.
To date, the structure of the nectary spur of Dendrobium finisterrae has not been studied in detail, and the present paper compares the structural organization of the floral nectary in this species with the spurs of other taxa. The nectary spur of D. finisterrae was examined by means of light microscopy (LM), scanning electron microscopy (SEM), and transmission electron microscopy (TEM). It is composed of a single layer of secretory epidermis and several layers of small and compactly arranged subepidermal secretory cells. The secretory cells have thick cellulosic cell walls with primary pits. The secretory tissue is supplied by vascular bundles that run beneath in ground parenchyma and are additionally surrounded by strands of sclerenchymatous fibers. The flowers of the investigated species displayed morphological features characteristic of bee-pollinated taxa, as they are zygomorphic, creamy-green coloured with evident nectar guides. They also emit a weak but nice scent. However, they possess some characters attributed to bird-pollinated flowers such as a short, massive nectary spur and collenchymatous secretory tissue that closely resembles the one found in the nectaries of certain species that are thought to be bird-pollinated. This similarity in anatomical organization of the nectary, regardless of geographical distribution and phylogeny, strongly indicates convergence and appears to be related to pollinator-driven selection.
Floral nectaries of Inula helenium L. only occurred in disc florets and were situated above the inferior ovary. The shape of the investigated glands (five-armed star with rounded tips and deep incisions - observed from above) clearly differed from the shape of the nectaries of other Asteraceae, also the height of nectary was much lower (129 µm). The glandular tissue of the nectaries of elecampane was composed of a single-layered epidermis and 5--9 layers of secretory cells. Nectar was released through modified stomata, mainly arranged in the top part of the gland. The secretory cells were characterised by granular cytoplasm and the presence of a large, often lobate, cell nucleus. In the cytosol, numerous amoeboid plastids, mitochondria, Golgi bodies and ribosomes were present. In small vacuoles, myelin-like structures, fibrous material and vesicles with the content of substances which can be secretion, were observed. The plastid stroma showed different electron density and the presence of internal tubules and plastoglobules. Vesicular extensions forming bright zones were visible between the membranes of the nuclear envelope. Adjacent to the plasmalemma, as well as between the plasmalemma and the cell wall, secretory vesicles occurred, indicating the granulocrine mechanism of nectar secretion.
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In carnivorous plants, two types of nectaries occur: extra- floral nectaries, generally associated with prey luring, and floral ones associated with pollination. Nectar produced by extra-floral nectaries not only attracts prey but may also be involved in trapping prey and plays a role in myrmecophily. The diversity of nectary structure in carnivorous plants reflects complicated evolutionary routes in this unique ecological group.
The structure of nectaries in the flowers of Myosotis sylvatica Hoffm. was studied by light microscopy, scanning electron microscopy and transmission electron microscopy. A nectariferous gland of irregular disc shape surrounds the bottom of the four-lobed ovary. From the upper side, both the nectary and the ovary are protected by ligulate folds of a widening corolla tube with epidermis outgrowths. Nectar is secreted by modified stomata situated only on the top and abaxial nectary surface adjoining the corolla tube. Stomata are irregularly distributed, forming aggregates of 2-3 each. On the longitudinal sections of the receptacle, the nectariferous tissue is distinguished by small cells, dense protoplast content, and intercellular spaces of different sizes. Branches of phloem bundles are observed at the base of the nectariferous tissue.
The conducted study related to the structure of the floral nectaries of Rhododendron japonicum (A. Gray) J. V. Suringar ex E. H. Wilson. The structure of the secretory epidermis of the nectaries was analysed by using scanning electron microscopy (SEM). Rhododendron japonicum develops the superior pistil with a 5-loculed ovary equipped in five ribs. The nectary gland is located in the lower part of the ovary. In the nectary regions located on the extension of the ribs of the ovary, stomata were very numerous. In the upper part of the nectary, stomata were arranged individually or in small clusters, whereas at its half-height they formed stomatal areas. The stomata were at different growth stages. They were arranged in different directions. The stomata developed on the nectary surface according to the mosaic pattern. The stomata from the lower situated part of the nectary had a different structure than those occurring in the upper half of the nectary. The stomata in the nectaries of Rh. japonicum belong to the actinocytic type. The cuticle layer in the upper part of the nectary was better developed and had a characteristic sculpture, whereas in the lower part it was smooth.
The studies on Asphodelus albus Miller flowers were conducted in the Botanical Garden of the UMCS in Lublin in the years 2004-2005. The flower nectaries location was determined in a stereoscopic microscope. The nectaring abundance was studied with a pipette method described by Jabłoński and Szklanowska (1979), while pollen efficiency determined after Warakomska's ether method (1972). Pollen viability was computed in a sample of 400 grains after acetocarmine staining. The following measurements of pollen grains were made: the length of polar axis (P), equatorial longitudinal axis (EL) and equatorial transverse axis (ET). In Asphodelus albus flowers, there are three nectary glands located in the ovary septa whose outlets are situated in the upper part of the ovary. The nectar secretion starts in a dehiscing bud and persists until the withering stage of perianth leaves. Considering the size of monocolpate pollen grains of Asphodelus albus, they are ranked among great, whereas their shape assumed flattened and circular at the polar view. In the Poland climatic conditions, a pollen showed high vitality (98%). The Asphodelus albus plants constitute a valuable source of nutrition for the pollinators as a single flower generated on average 4,22 mg sugars and 0,2 mg of pollen grains.
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