Nectar and pollination drops: how different are they?

Background

Pollination drops and nectars (floral nectars) are secretions related to plant reproduction. The pollination drop is the landing site for the majority of gymnosperm pollen, whereas nectar of angiosperm flowers represents a common nutritional resource for a large variety of pollinators. Extrafloral nectars also are known from all vascular plants, although among the gymnosperms they are restricted to the Gnetales. Extrafloral nectars are not generally involved in reproduction but serve as ‘reward’ for ants defending plants against herbivores (indirect defence).

Scope

Although very different in their task, nectars and pollination drops share some features, e.g. basic chemical composition and eventual consumption by animals. This has led some authors to call these secretions collectively nectar. Modern techniques that permit chemical analysis and protein characterization have very recently added important information about these sugary secretions that appear to be much more than a ‘reward’ for pollinating (floral nectar) and defending animals (extrafloral nectar) or a landing site for pollen (pollination drop).

Conclusions

Nectar and pollination drops contain sugars as the main components, but the total concentration and the relative proportions are different. They also contain amino acids, of which proline is frequently the most abundant. Proteomic studies have revealed the presence of common functional classes of proteins such as invertases and defence-related proteins in nectar (floral and extrafloral) and pollination drops. Invertases allow for dynamic rearrangement of sugar composition following secretion. Defence-related proteins provide protection from invasion by fungi and bacteria. Currently, only few species have been studied in any depth. The chemical composition of the pollination drop must be investigated in a larger number of species if eventual phylogenetic relationships are to be revealed. Much more information can be provided from further proteomic studies of both nectar and pollination drop that will contribute to the study of plant reproduction and evolution.Key words: Nectar, pollination drop, ovular secretion, plant reproduction, proteins, sugars, gymnosperms, angiosperms, plant–animal interaction     

Nectar robbing in Ipomopsis aggregata : effects on pollinator behavior and plant fitness

Hummingbirds foraging in alpine meadows of central Colorado, United States, face a heterogeneous distribution of nectar rewards. This study investigated how variability in nectar resources caused by nectar-robbing bumblebees affected the foraging behavior of hummingbird pollinators and, subsequently, the reproductive success of a host plant (Ipomopsis aggregata). We presented hummingbirds with experimental arrays of I. aggregata and measured hummingbird foraging behavior as a function of known levels of nectar robbing. Hummingbirds visited significantly fewer plants with heavy nectar robbing (over 80% of available flowers robbed) and visited fewer flowers on those plants. These changes in hummingbird foraging behavior resulted in decreased percent fruit set as well as decreased total seed set in heavily robbed plants. These results indicate that hummingbird avoidance of nectar-robbed plants and flowers reduces plant fitness components. In addition, our results suggest that the mutualisms between pollinators and host plants may be affected by other species, such as nectar robbers. Received: 22 April 1998 / Accepted: 12 May 1998     

Nectar thieves or invited pollinators? A case study of tansy flowers and common house mosquitoes

Mosquitoes are generally considered nectar thieves that do not contribute to the pollination of the flowers they visit. Here we tested the hypothesis that the common house mosquito, Culex pipiens, contributes to the pollination of tansies, Tanacetum vulgare, and possibly the pollination of other members of the Asteraceae (Achillea millefolium, Leucanthemum vulgare, Solidago canadensis). To test this hypothesis, we (1) field-collected mosquitoes probing inflorescences of T. vulgare, A. millefolium, and L. vulgare, and recorded the number and distribution of pollen grains on their bodies, (2) exposed laboratory-reared Cx. pipiens to inflorescences of T. vulgare, A. millefolium, and S. canadensis, and (3) ran pollination experiments with Cx. pipiens and T. vulgare in a greenhouse. We found (1) that 41 of 164 field-collected Cx. pipiens carried pollen, (2) that 48, 34, and 34 % of Cx. pipiens accumulated pollen from T. vulgare, A. millefolium, and S. canadensis, respectively, during floral visits of greenhouse-grown specimens, and (3) that cross-pollination by Cx. pipiens resulted in significant seed set of T. vulgare in pollination experiments. Based on our observations that Cx. pipiens are frequent floral visitors and are able to carry pollen and to induce seed set in T. vulgare, it is clear that Cx. pipiens plays a role in the pollination of T. vulgare, and possibly other members of the Asteraceae.     

Floral Biology and Pollination Mechanisms in Two Viola Species—From Nectar to Pollen Flowers?

The genus Viola is represented by four related species in Brazil belonging to section Leptidium, one of the most primitive sections in the genus. Floral biology and pollination by bees were studied in Viola cerasifolia and V. subdimidiata in high-altitude areas in south-eastern Brazil. Flowers are zygomorphic and spurred. The five stamens are arranged in a cuff around the ovary, and pollen is released by means of apical connective projections, which form a cone surrounding the base of the style. The connective projections of the inferior stamens are elongated and curved to form a hook-shaped structure. Nectar-secreting tissue can occur in the basal connective appendages of the inferior stamens, which project into the spur. Flowers of V. subdimidiata secreted a mean volume of 0.14 micro l nectar over a 24-h period; approx. 40 % of flowers did not secrete any nectar. The main pollinators of these Viola species are female bees belonging to the genus Anthrenoides (Andrenidae), which search mainly for pollen. These bees seem to be oligolectic and obtain large amounts of pollen from Viola by vibrating the flowers or by moving the hook repeatedly back and forth. Males of Anthrenoides patrol Viola clusters and also feed on nectar, acting as secondary pollinators. The basic floral structure in the genus Viola fits that of 'nectar flowers'. The uncommon hook-shaped projections, scanty nectar production, and behaviour of pollinators suggest that V. cerasifolia and V. subdimidiata are shifting their reward for pollinators from nectar to pollen. Based on floral morphology, this shift may be widespread in Viola sect. Leptidium.     

Do Honeybees Shape the Bacterial Community Composition in Floral Nectar?

Floral nectar is considered the most important reward animal-pollinated plants offer to attract pollinators. Here we explore whether honeybees, which act as pollinators, affect the composition of bacterial communities in the nectar. Nectar and honeybees were sampled from two plant species: Amygdalus communis and Citrus paradisi. To prevent the contact of nectar with pollinators, C. paradisi flowers were covered with net bags before blooming (covered flowers). Comparative analysis of bacterial communities in the nectar and on the honeybees was performed by the 454-pyrosequencing technique. No significant differences were found among bacterial communities in honeybees captured on the two different plant species. This resemblance may be due to the presence of dominant bacterial OTUs, closely related to the Arsenophonus genus. The bacterial communities of the nectar from the covered and uncovered C. paradisi flowers differed significantly; the bacterial communities on the honeybees differed significantly from those in the covered flowers’ nectar, but not from those in the uncovered flowers’ nectar. We conclude that the honeybees may introduce bacteria into the nectar and/or may be contaminated by bacteria introduced into the nectar by other sources such as other pollinators and nectar thieves.     

Are Nectar Sugar Composition and Corolla Tube Length Related to the Diversity of Insects that Visit Asteraceae Flowers?

Abstract: In this work, we analysed interspecific variation in nectar sugar composition, corolla tube length, and the diversity of floral visitors of 35 Asteraceae species. The potential correlations between these variables could arise either as a result of selection to improve pollinator attractiveness or simply as a consequence of phylogenetic constraints. Samples of nectar and flowers, and data on floral visitors, were obtained from living plants in natural populations from Argentina. Asteraceae species showed a large variability in corolla tube length. Nectar of most species presented a larger proportion of hexoses than sucrose. All species were visited by numerous insects belonging to 2 different orders. Results showed that floral traits are not significantly correlated with the diversity of floral visitors. These characters seem to be linked to the phylogeny of the species. Early branching species (species phylogenetically close to the root of the Asteraceae tree) tend to have longer corollas, higher sucrose proportions and lesser diversity of floral visitors than late branching species. Considering that longer corolla tubes and higher nectar sucrose percentages may indicate some specialization in the pollination system, we suggest that there is an evolutionary tendency toward generalist pollination systems within the family.     

Raising the Sugar Content – Orchid Bees Overcome the Constraints of Suction Feeding through Manipulation of Nectar and Pollen Provisions

Unlike most other bees, the long-tongued orchid bees ingest nectar using suction feeding. Although long tongues allow exploitation of flowers with deep spurs, the energy intake rate is optimal at 10–20% lower nectar sugar concentrations compared to that of lapping bees. This constraint might be compensated by a higher digestive throughput. Additionally, orchid bees might evaporate water from regurgitated droplets of crop contents. We found male Euglossa championi (n = 10) and Euglossa dodsoni (n = 12) to regularly regurgitate droplets of crop content to the base of their proboscis, generating a fluid film between the proximal parts of the galeae, glossa and labial palps. Rhythmic movements of the proboscis may help to increase convection. There was a significant change in sugar concentration between the initially imbibed solution and the resulting crop content (P<0.05) and the time individual bees had engaged in this liquid exposure behavior was positively correlated with the resulting crop sugar concentration. Female Euglossa townsendi and Euglossa viridissima showed the same behavior. Additionally, they manipulated their nectar-enriched pollen provisions for extensive periods of time before deposition in brood cells. The deposited pollen loads (n = 14) showed a significantly higher sugar concentration than the sugar-water available to the bees (P<0.001). Thus, both male and female euglossines show behaviors that promote evaporative water loss from nectar. We suggest that the behaviors have evolved in concert with suction feeding on dilute nectar from deep floral tubes.     

Nectar and Pollen Sources for Honeybee （Apis cerana cerana Fabr.） in Qinglan Mangrove Area, Hainan Island, China

In the present study, nectar and pollen sources for honeybee （Apls cerana cerana Fabr.） were studied in Qlnglan mangrove area, Hainan Island, China, based on microscopic analysis of honey and pollen load （corblcular and gut contents） from honeybees collected In October and November 2004. Qualitative and quantitative melittopalynologlcal analysis of the natural honey sample showed that the honey is of unlfloral type with Mimosa pudlca L. （Mlmosaceae） as the predominant （89.14%） source of nectar and pollen for A. cerana cerana In October. Members of Araceae are an Important minor （3%-15%） pollen type, whereas those of Arecaceae are a minor （〈3%） pollen type. Pollen grains of Nypa fruticans Wurmb., Rhlzophora spp., Excoecarla agallocha L., Lumnitzera spp., Brugulera spp., Kandella candel Druce, and Ceriops tagal （Perr.） C. B. Rob. are among the notable mangrove texa growing In Qinglan mangrove area recorded as minor taxa In the honey. The absolute pollen count （I.e. the number of pollen grains/10 g honey sample） suggests that the honey belongs to Group V （〉1 000 000）. Pollen analysis from the corblcular and gut contents of A. cerana cerana revealed the highest representation （95.60%） of members of Sonneratia spp. （Sonneratlaceae）, followed by Bruguiera spp. （Rhizophoraceae）, Euphorblaceae, Poaceae, Fabaceae, Arecaceae, Araceae, Anacardlaceae, and Rublaceae. Of these plants, those belonging to Sonneratla plants are the most Important nectar and pollen sources for A. cerana cerana and are frequently foraged and pollinated by these bees in November.     

Nectar bacteria,but not yeast,weaken a plant–pollinator mutualism

Mutualistic interactions are often subject to exploitation by species that are not directly involved in the mutualism. Understanding which organisms act as such ‘third-party’ species and how they do so is a major challenge in the current study of mutualistic interactions. Here, we show that even species that appear ecologically similar can have contrasting effects as third-party species. We experimentally compared the effects of nectar-inhabiting bacteria and yeasts on the strength of a mutualism between a hummingbird-pollinated shrub, Mimulus aurantiacus, and its pollinators. We found that the common bacterium Gluconobacter sp., but not the common yeast Metschnikowia reukaufii, reduced pollination success, seed set and nectar consumption by pollinators, thereby weakening the plant–pollinator mutualism. We also found that the bacteria reduced nectar pH and total sugar concentration more greatly than the yeasts did and that the bacteria decreased glucose concentration and increased fructose concentration whereas the yeasts affected neither. These distinct changes to nectar chemistry may underlie the microbes'' contrasting effects on the mutualism. Our results suggest that it is necessary to understand the determinants of microbial species composition in nectar and their differential modification of floral rewards to explain the mutual benefits that plants and pollinators gain from each other.     

The Sweet Tooth of Adult Parasitoid Cotesia rubecula: Ignoring Hosts for Nectar?

Investing time and energy into survival and reproduction often presents a trade-off to many species of animals. In parasitic wasps, both hosts and sugar sources contribute to the forager's fitness but are often found in different locations. The decision to search for hosts or for food can have a strong impact on fitness when the forager's lifetime is short and resources are not abundant. We investigated the tendency of flowers and hosts to attract 1-day-old female Cotesia rubecula Marshall (Hymenoptera: Braconidae) with different feeding histories in a wind tunnel. Only well-fed wasps exhibited a preference for hosts. In comparison, unfed wasps visited hosts and flowers in equal proportions. Feeding experience had a strong impact on the searching behavior and the number of landings on both resources. Host and food stimuli seem to be equally attractive to hungry parasitic wasps such as C. rubecula. We expect that under field conditions the time available for active food searching in female C. rubecula is short and influenced by the presence of hosts.     

Nectar to improve parasitoid fitness in biological control: Does the sucrose:hexose ratio matter?

The consumption of saccharide-rich foods such as floral nectar is crucial for the survival of many adult parasitoid wasps. The importance to parasitoids of nectar quality, with regards to its sucrose:hexose ratio, was investigated. Nectar, an aqueous solution of sugars, amino acids and other compounds, differs between plant species. Nectar composition is dominated by sucrose, glucose and fructose. Previous studies have shown that the ratio of sucrose to hexose (glucose+fructose) sugars can explain nectar associations in a range of flower visiting arthropods. It has been suggested that this ratio may be important in terms of parasitoid fitness. Analysis of floral nectar from fourteen plant species confirmed that the sucrose/hexose ratio significantly differed between species. An opportunity to select floral resources based on this measure of nectar quality arose and highlighted the potential to utilize native flowering plant species in place of the seven most commonly deployed, which are usually not native to the countries in which they are used.Results presented in this paper indicate, however, that the sucrose/hexose ratio is not a significant factor explaining parasitoid longevity. The hymenopteran parasitoids Diadegma semiclausum (Ichneumonidae) and Dolichogenidea tasmanica (Braconidae) were fed 40% w/w sugar solutions, differing in their sugar ratios. Solutions were classified as either sucrose-dominant (ratio >0.99), sucrose-rich (ratio 0.5–0.99), hexose-rich (ratio 0.1–0.499) or hexose-dominant (ratio <0.1). No significant differences in parasitoid longevity were found between the different treatments for either species. This suggests there is not an optimal sucrose/hexose ratio for parasitoid wasps, although a greater number of parasitoid species should ideally be tested to confirm if this is true for the wider parasitoid taxonomic groups.     

Why Do Males of Parnassius imperator Fight for Bare Rocks but Not the Nectar Flower During Mate Selection?

Territoriality is an important male behavioral trait used in mate selection of butterflies, but the factors determining the outcome of territorial contests remain controversial. We studied the foraging and territorial behaviors of males in the Parnassius imperator butterfly. Adult males were selective about nectar sources, but they did not defend nectar flowers. By contrast, they fought over areas of bare rocks because bare rocks were used as lek locations to attract and court receptive females. The ownership of a lek, rather than sex pheromones or wing pigments, played the key role in attracting females. In an owner replacement experiment, the owner of a territory always defeated intruders, indicating that perching order, not relative body size, determined the outcome of territorial contests. Territories are small areas of bare rocks that lack food resources, so intruders do not challenge the ownership of a resident in a location where they can easily find vacant territories. Males that first perch in a territory have an advantage compared to intruders in that they can forage on flowers in the vicinity of the territory. This may help explain why satiated owners always defeated fatigued intruders. Our findings suggest that early perching on bare rock allows male butterflies of P. imperator to occupy a territory, attract receptive females and exclude later‐arriving competitors.     

Nectar intake of white-bellied sunbirds (Cinnyris talatala): can meal size be inferred from feeding duration?

A positive relationship between feeding duration and meal size of avian nectarivores has often been assumed in earlier studies. We investigated whether feeding duration can be used as a surrogate for the amount of sugar solution ingested by white-bellied sunbirds, Cinnyris (Nectarinia) talatala. Feeding durations of sunbirds consuming three sucrose concentrations (10%, 20%, and 40% w/w) were measured using an infrared photodetection system, and the amounts consumed were recorded simultaneously by weighing the feeder throughout the experiment. For all three diet concentrations, a positive relationship was found between the time spent feeding per 30 min and the mass consumed. Therefore, feeding duration is demonstrated to be an index of the amount ingested on a particular sugar concentration. The rate of ingestion, however, depended on the sugar concentration, with the highest rate at the lowest concentration of 10% and the lowest rate at the 40% concentration. Less total time was spent feeding on the 20% solution than on the 10% solution, but time increased on the 40% diet because of viscosity effects. There appeared to be a weak relationship between feeding patterns and sex, but this was not significant, probably because of interindividual variation.     

Syntheses and spectroscopic properties of α- and β-phosphatidylcholines and phosphatidylethanolamines

The widely used partial synthesis of phospholipids via deacylation of naturally occurring phospholipids, followed by reacylation with fatty acid anhydrides, is accompanied by phosphoryl migration. The resulting mixture of α- and β-phospholipids was separated by short-column chromatography. Milder acylation procedures in which no phosphoryl migration occurs, were developed. 1,2-Dilinoleoyl-sn-glycero-3-phosphocholine was prepared in 50% yield by acylation of sn-glycero-3-phosphocholine (GPC) with N-linoleoylimidazole. Detailed NMR and infrared spectra of α- and β-phosphatidylcholines (PCs) and -ethanolamines (PEs) are reported and the differences between isomers discussed.     

Phenotypic plasticity and longevity in plants and animals: cause and effect?

Immobile plants and immobile modular animals outlive unitary animals. This paper discusses competing but not necessarily mutually exclusive theories to explain this extreme longevity, especially from the perspective of phenotypic plasticity. Stem cell immortality, vascular autonomy, and epicormic branching are some important features of the phenotypic plasticity of plants that contribute to their longevity. Monocarpy versus polycarpy can also influence the kind of senescent processes experienced by plants. How density-dependent phenomena affecting the establishment of juveniles in these immobile organisms can influence the evolution of senescence, and consequently longevity, is reviewed and discussed. Whether climate change scenarios will favour long-lived or short-lived organisms, with their attendant levels of plasticity, is also presented.     

Dogmatisms and Catechisms — Ethics and Companion Animals

Abstract

The current reconsideration of the place in nature of human beings unfortunately continues to be an acrimonious one. All too often the debate is more akin to a warlike encounter where each side attempts to gain control or the upper hand than a search for points of agreement. Given this context, it is important to entertain views that emanate from different cultural traditions as a way to infuse the debate with new life. Students of Native American culture have consistently pointed out that the essential concepts of life balance and reciprocity represented there may serve as useful points of consideration as we struggle with the appropriate relationships with animals and nature. This article presents a representative Zuni story, told by Governor Robert E. Lewis, that illustrates these notions.     

DCCP and DICP： Construction and Analyses of Databases for Copper- and Iron-Chelating Proteins

Copper and iron play important roles in a variety of biological processes, especially when being chelated with proteins. The proteins involved in the metal binding, transporting and metabolism have aroused much interest. To facilitate the study on this topic, we constructed two databases （DCCP and DICP） containing the known copper- and iron-chelating proteins~ which are freely available from the website http：//sdbi.sdut.edu.cn/en. Users can conveniently search and browse all of the entries in the databases. Based on the two databases, bioinformatic analyses were performed, which provided some novel insights into metalloproteins.     

Biosynthesis of geraniol and nerol and their β-d-glucosides in Perlargonium graveolens and Rosa dilecta

1. 3R-[2-(14)C]Mevalonate was incorporated into geranyl and neryl beta-d-glucosides in petals of Rosa dilecta in up to 10.6% yield, and the terpenoid part was specifically and equivalently labelled in the moieties derived from isopentenyl pyrophosphate and 3,3-dimethylallyl pyrophosphate. A similar labelling pattern, with incorporations of 0.06-0.1% was found for geraniol or nerol formed in leaves of Pelargonium graveolens The former results provide the best available evidence for the mevalonoid route to regular monoterpenes in higher plants. 2. Incorporation studies with 3RS-[2-(14)C,(4R)-4-(3)H(1)]-mevalonate and its (4S)-isomer showed that the pro-4R hydrogen atom of the precursor was retained and the pro-4S hydrogen atom was eliminated in both alcohols and both glucosides. These results suggest that the correlation of retention of the pro-4S hydrogen atom of mevalonate with formation of a cis-substituted double bond, such as has been found in certain higher terpenoids, does not apply to the biosynthesis of monoterpenes. It is proposed that either nerol is derived from isomerization of geraniol or the two alcohols are directly formed by different prenyltransferases. Possible mechanisms for these processes are discussed. 3. The experiments with [(14)C,(3)H]mevalonate also show that in these higher plants, as has been previously found in animal tissue and yeast, the pro-4S hydrogen atom of mevalonate was lost in the conversion of isopentenyl pyrophosphate into 3,3-dimethylallyl pyrophosphate.     

Zinc and diabetes — clinical links and molecular mechanisms

Zinc is an essential trace element crucial for the function of more than 300 enzymes and it is important for cellular processes like cell division and apoptosis. Hence, the concentration of zinc in the human body is tightly regulated and disturbances of zinc homeostasis have been associated with several diseases including diabetes mellitus, a disease characterized by high blood glucose concentrations as a consequence of decreased secretion or action of insulin. Zinc supplementation of animals and humans has been shown to ameliorate glycemic control in type 1 and 2 diabetes, the two major forms of diabetes mellitus, but the underlying molecular mechanisms have only slowly been elucidated. Zinc seems to exert insulin-like effects by supporting the signal transduction of insulin and by reducing the production of cytokines, which lead to beta-cell death during the inflammatory process in the pancreas in the course of the disease. Furthermore, zinc might play a role in the development of diabetes, since genetic polymorphisms in the gene of zinc transporter 8 and in metallothionein (MT)-encoding genes could be demonstrated to be associated with type 2 diabetes mellitus. The fact that antibodies against this zinc transporter have been detected in type 1 diabetic patients offers new diagnostic possibilities. This article reviews the influence of zinc on the diabetic state including the molecular mechanisms, the role of the zinc transporter 8 and MT for diabetes development and the resulting diagnostic and therapeutic options.