Nectar production and transportation in the nectaries of the female Cucumis sativus L. flower during anthesis

Summary. In an effort to gain a greater understanding of nectar production, we studied the dynamic mechanisms of starch accumulation and transformation and nectar transportation in the Cucumis sativus L. female flower. Starch, which is the main precursor of nectar, accumulates in the epidermis and underlying parenchyma, with the most active accumulation occurring in the parenchyma cells within 3 days prior to anthesis. Thereafter, the starch was successively hydrolyzed and the hydrolyte was transported from the amyloplasts to vacuoles, suggesting that amyloplasts and vacuoles are the centers of nectar production. In addition, we observed few plasmodesmata and the presence of invaginated plasmalemma and electron-dense material in the intercellular spaces, suggesting that the apoplast system is involved in nectar transportation in an ATPase-dependent fashion.Correspondence and reprints: College of Life Sciences, Peking University, Beijing, 100871, Peoples Republic of China     

Floral nectaries, nectar production dynamics and chemical composition in six ipomoea species (convolvulaceae) in relation to pollinators

BACKGROUND AND AIMS: Floral nectaries and nectar features were compared between six Argentinian Ipomoea species with differences in their pollinator guilds: I. alba, I. rubriflora, I. cairica, I. hieronymi var. hieronymi, I. indica, and I. purpurea. METHODS: Pollinators were recorded in natural populations. The morpho-anatomical study was carried out through scanning electron and light microscopy. Nectar sugars were identified via gas chromatography. Nectar production and the effect of its removal on total nectar sugar amount were determined by using sets of bagged flowers. KEY RESULTS: Hymenopterans were visitors of most species, while hummingbirds visited I. rubriflora and sphingids I. alba. All the species had a vascularized discoidal nectary surrounding the ovary base with numerous open stomata with a species-specific distribution. All nectar samples contained amino acids and sugars. Most species had sucrose-dominant nectars. Flowers lasted a few hours. Mean nectar sugar concentration throughout the lifetime of the flower ranged from 34.28 to 39.42 %, except for I. cairica (49.25 %) and I. rubriflora (25.18 %). Ipomoea alba had the highest nectar volume secreted per flower (50.12 microL), while in the other taxa it ranged from 2.42 to 12.00 microL. Nectar secretion began as soon as the flowers opened and lasted for a few hours (in I. purpurea, I. rubriflora) or it was continuous during the lifetime of the flower (in the remaining species). There was an increase of total sugar production after removals in I. cairica, I. indica and I. purpurea, whereas in I. alba and I. rubriflora removals had no effect, and in I. hieronymi there was a decrease in total sugar production. CONCLUSIONS: The chemical composition, production dynamics and removal effects of nectar could not be related to the pollinator guild of these species. Flower length was correlated with nectary size and total volume of nectar secreted, suggesting that structural constraints may play a major role in the determination of nectar traits of these species.     

Floral ontogeny of Cneorum tricoccon L. (Rutaceae)

The floral ontogeny of the Spurge olive (Cneorum tricoccon L.) is studied by means of scanning electron microscopic observations. Special attention is paid to the sequence of initiation of the floral parts, the occurrence of septal cavities, and the development of the nectariferous tissue. The nectary disc arises as a receptacular outgrowth below the ovary and independently from stamen development. By the extensive growth of this voluminous androgynophore, stamen filaments become enclosed by nectary tissue and as a result, they are seated in pits between the lobes of the disc. Between ovary and style, three lobes are present, which are covered with stomata – their function is unknown. The significance of the unusual trimery of the flower is discussed. Floral developmental evidence supports a Rutalean affinity, although more ontogenetic investigations are needed in Rutaceae, subfamily Spathelioideae.     

Time‐course proteome analysis of developing extrafloral nectaries of Ricinus communis

Floral and extrafloral nectaries are unique organs that secrete energy rich chemical components, but their contribution for nectar production is largely unknown. Here, we present the first comparative proteome dataset of four developmental stages of the extrafloral nectaries from castor plant (Ricinus communis), an important biofuel crop. Respectively, from stage I—IV, we identified 626, 613, 449 and 356 proteins, respectively, summing up 882 nonredundant proteins. Surprisingly, we identified two isoforms of the potent toxin ricin, indicating that ricin expression is not limited to seeds, but it may serve a general defense purpose for the castor plant. To date, this is the most complete dataset of proteins either from floral or extrafloral nectaries, thus contributing to lay the foundations for investigations on their ecological and evolutionary importance.     

Enhanced UV-B radiation,flower attributes and pollinator behaviour in Cistus creticus: a Mediterranean field study

The aim of this investigation was to examine the reasons for the higher pollination success in Cistus creticus under enhanced UV-B radiation (Stephanou & Manetas 1998). Thus, a selected array of floral attributes as well as the frequency and duration of insect visits were studied in the field under ambient or ambient plus supplemental UV-B radiation, simulating a 15% ozone depletion over Patras (38.3° N, 29.1° E). Video-recording revealed two categories of visitors, i.e. true pollinators (bees) and nectar thieves. The frequency of visits to both control and UV-B treated plants was the same and independent of whether the UV-B tubes were on or off during video recording. UV-B radiation had no effect on gross floral morphology (petal surface area, number of pollen grains, stamens and ovules, optical properties of petals and stamens), yet nectary size was almost doubled. In addition, the duration of insect visits was significantly longer on UV-B treated plants, provided that the UV-B tubes were off during monitoring. The differences were abolished during the part of the day that the tubes were on, indicating that the insects were annoyed by supplemental UV-B radiation. These results are consistent with the nectaries producing larger quantities of nectar, which caused the insects to stay longer on flowers of UV-B treated plants and improved pollination success.     

Nectar-carbohydrate production and composition vary in relation to nectary anatomy and location within individual flowers of several species of Brassicaceae

Nectar-carbohydrate production and composition were investigated by high-performance liquid chromatography and enzymology in nine species from five tribes of the Brassicaceae. In six species (Arabidopsis thaliana (L.) Heynh., Brassica napus L., B. rapa L., Lobularia maritima (L.) Desv., Raphanus sativus L., Sinapis arvensis L.) that produced nectar from both lateral nectaries (associated with the short stamens) and median nectaries (outside the long stamens), on average 95% of the total nectar carbohydrate was collected from the lateral ones. Nectar from these glands possessed a higher glucose/fructose ratio (usually 1.0–1.2) than that from the median nectaries (0.2–0.9) within the same flower. Comparatively little sucrose was detected in any nectar samples except from Matthiola bicornus (Sibth. et Sm.) DC., which possessed lateral nectaries only and produced a sucrose-dominant exudate. The anatomy of the nectarial tissue in nectar-secreting flowers of six species, Hesperis matronalis L., L. maritima, M. bicornus, R. sativus, S. arvensis, and Sisymbrium loeselii L., was studied by light and scanning-electron microscopy. Phloem alone supplied the nectaries. However, in accordance with their overall nectar-carbohydrate production, the lateral glands received relatively rich quantities of phloem that penetrated far into the glandular tissue, whereas median glands were supplied with phloem that often barely innervated them. All nectarial tissue possessed modified stomata (with the exception of the median glands of S. loeselii, which did not produce nectar); further evidence was gathered to indicate that these structures do not regulate nectar flow by guard-cell movements. The numbers of modified stomata per gland showed no relation to nectar-carbohydrate production. Taken together, the data on nectar biochemistry and nectary anatomy indicate the existence of two distinct nectary types in those Brassicacean species that possess both lateral and median nectaries, regardless of whether nectarial tissue is united around the entire receptacle or not. It is proposed that the term “nectarium” be used to represent collectively the multiple nectaries that can be found in individual flowers. Received: 21 July 1997 / Accepted: 19 September 1997     

Nectar secretion inAbutilon: a new model

Summary Nectary trichomes ofAbutilon striatum secrete copious amounts of sucrose, fructose and glucose. The nectar emerges from transient pores in the cuticle overlying the trichome tip cells. Calculations of the required transmembrane fluxes, either across the tip cell plasmalemma or across the cell membrane of the whole trichome, give very high rates compared with those obtained from other situations in plants and, therefore, cast doubt on the possibility that nectar secretion inAbutilon is an eccrine process. Quantitative evaluation of the possibility of granulocrine secretion, by successive fusion of vesicles with the cell membrane, suggests that this is an even less probable mechanism of secretion. Rapid freezing followed by freeze-substitution or freeze-fracture replication reveals that an extensive secretory reticulum (SR) is present within the hair cells. As similar micrographs are obtained from conventional, chemical fixation it is argued that the secretory reticulum is a relatively stable endomembrane system. Freeze-fracture and freeze-substitution micrographs show that this internal membrane system is closely associated with the plasmalemma. Taken together with other structural information, as well as physiological data, it is concluded that prenectar is actively loaded into the secretory reticulum of all trichome cells. Increase in hydrostatic pressure within this compartment leads to the opening of sphincters which connect the cisternal space of the SR to the outside of the plasmalemma. Thus a pulse of nectar is forcibly expelled into an apoplastic compartment sealed to the outside by the impermeable cuticle and on the inside by the plasmalemma. As this apoplastic compartment is also sealed at the stalk cell, the only route for pressure release is via the transient pores which overlay the tip cell. Distension renders these patent so that, again, pulsed secretion is observed. This hypothesis overcomes the necessity for envisaging excessively high transmembrane fluxes or rates of vesicle fusion. It would imply the need for a continuing supply of prenectar to the hair cells accompanied by active loading into the SR. This loading process may well be supported by the hydrolysis of sucrose to glucose and fructose and is probably the site where ions and other low molecular weight solutes are filtered from the nectar.     

Floral color changes in Boswellia sacra Flueck. (Burseraceae): A dialogue between plant and pollinator

The present study provides new information about the reproductive biology of Boswellia sacra (Burseraceae), focusing on the nectary and its attractiveness for pollinators. The nectary disc changes its color from yellow to orange and red during the flower development. The colors are related to the main period of the stigmatic receptivity, to the dehiscence of anthers with pollen presentation and the nectar secretion. Pollinators preferentially visit the flowers in the “yellow” phase and neglect the “red phase”. This suggests a sophisticated dialogue between the plant and its pollinators. The color change from yellow to red occurs in a very short time (less than 24 h) and it is due to the accumulation of anthocyanins. Despite this dialogue between plant and pollinators, the number of fruits is often scanty.     

Nectar, nectaries, flower visitors, and breeding system in five terrestrial Orchidaceae from central Argentina

Floral nectar sugar composition, nectary anatomy, and visitors are studied in five Argentine Orchidaceae, from 18 populations. Hand-pollinations were performed to evaluate their breeding system. We found two different types of perigonal nectaries located either in the spur (Habenaria gouriieana, H. hieronymi, Habenariinae), or in the basal lateral parts of the labellum (Beadlea dutraei, Pelexia bonariensis, Stenorrhynchos orchioides, Spiranthinae). The spur ofHabenaria is a nonvascularised and nonstructural nectary. The inner epidermis bears one-celled long papillae. In bud stage, the papillae are filled with starch grains, but when the flower opens and nectar secretion starts, they show no starch grains. This fact may indicate that starch is a source for some of the secreted nectar. In the remainder genera, the lateral basal parts of the labellum are secretory. The two glands are located in the adaxial basal lateral faces of the labellum. These nectaries are structural and nonvascularised.Stenorrhynchos produces abundant, concentrated nectar (40–50%).Habenaria gourlieana accumulates copious nectar in a lower concentration (<20%), whereas the other species produce small quantities of concentrated nectar (ca. 50%). Three of the studied species have sucrose predominant nectar (Beadlea dutrael, Habenaria gourlieana, andPelexia bonariensis) whileH. hieronymi, Stenorrhynchos orchioides have hexose predominant ones. Nectar removal and/or pollination induce flower senescence.H. gouriieana is visited by sphingids,S. orchioides by hummingbirds, andB. dutrael by bees. For the two other species we did not record flower visitors.Pelexia bonariensis, B. dutrael, andS. orchiodes are self-compatible species but a pollinator is needed.