A model for nectar secretion in animal-pollinated plants

Summary A game theoretic model was developed for nectar secretion in animal-pollinated plants in order to examine how the total amount of resources allocated to flowers affects the spread of nectarless plants. It was assumed that pollinators concentrate on patches whose nectar rewards are relatively large compared to other patches and if pollinators visit a patch, they concentrate on the plants whose nectar rewards are relatively large compared to other plants in the patch. It was shown that plants are more likely to secrete nectar in populations where the total amount of resources allocated to flowers is large. It was also shown that strong interplant competition, strong interpatch competition and the nectar discrimination of the pollinators are also important factors for nectar secretion. However, if the total amount of resources allocated to flowers is sufficiently large, plants would secrete nectar even if competition is not very strong and nectar discrimination is not so precise.     

Dark purple nectar as a foraging signal in a bird-pollinated Himalayan plant

? Some plants secrete coloured nectar to attract pollinators, but little is known about the chemical origins of nectar colouration and its ecological function. Leucosceptrum canum stands out as the only plant with coloured nectar recorded in the Himalayas. Here, we focused on the compound associated with the dark colour of the nectar, as well as its secretion dynamics during the flowering season and its relationship to pollinators. ? Fresh nectar was analysed by semi-preparative reversed-phase high-performance liquid chromatography (HPLC), LC-MS and HRESIMS (high resolution electronspray ionization mass spectroscopy) to determine which compound causes the nectar colouration. Behavioural experiments were conducted with birds and honeybees to elucidate the effect of the nectar colour and volume on pollinators. ? We identified a purple anthocyanidin, 5-hydroxyflavylium, as a natural nectar product for the first time. Two short-billed birds were found to pollinate this plant, which employs two nectar-based mechanisms to direct bird pollinators to reproductively active flowers, controlling nectar palatability and presenting a foraging signal for birds by altering nectar volume and colour in a developmental stage-specific manner. ? 5-Hydroxyflavylium was found to be the cause of the nectar colouration, the function of which is to act as a foraging signal to increase pollination efficiency through nectar visibility and palatability.     

FLORAL BIOLOGY OF NEPENTHES GRACILIS (NEPENTHACEAE) IN SUMATRA

Nepenthes gracilis, a dioecious carnivorous plant, has inconspicuous flowers lacking petals. Nectaries distributed on the upper surface of four sepals secrete dilute nectar (3%–12% sugar concentration) at night, but the nectar immediately disappears during the day by evaporation in the sunny environment of Sumatra. Male flowers have a higher nectar production rate but lower sugar concentration of nectar than female flowers. Flowers of both sexes were visited by pyralid moths at night and by calliphorid flies in the evening. Pollen was found attached on these insects visiting Nepenthes flowers. The pattern of nectar production of sepals is regarded as attracting nocturnal flying insects and avoiding ants, while the pitchers attract ants by nectar secreted on the pitcher rim.     

FUNCTION OF MENTZELIA NUDA (LOASACEAE) POSTFLORAL NECTARIES IN SEED DEFENSE

Mentzelia nuda is a coarse, short-lived perennial of the High Plains of North America. The flowers secrete nectar which serves as an attractant for pollinating bees. Postfloral nectar secretion attracts ants whose presence significantly enhances seed set. This is the first demonstration of advantage to postflowering activity of a floral nectary.     

The reproductive biology of Sophora fernandeziana (Leguminosae), a vulnerable endemic species from Isla Robinson Crusoe

Sophora fernandeziana is the only legume endemic to Isla Robinson Crusoe (Archipelago Juan Fernández, Chile); it is uncommon and becoming rare. Although its preservation status is listed as "vulnerable," as with many species, little is known of its reproductive biology. Flowering phenology, floral morphology, nectar features, breeding system, and visitors were analyzed in two populations. Flowering is from late winter to early spring. Flowers last 6 d and have a number of ornithophilous features. A floral nectary begins to secrete highly concentrated nectar 48 h after flowers open. Nectar secretion increases as the flower ages but culminates in active nectar reabsorption as the flower senesces. Nectar production is negatively affected by nectar removal. Self-pollen germinates and tubes grow down the style. However, pollen tubes were only observed to enter the ovaries in open pollinated styles, suggesting the possibility of an ovarian self-incompatibility mechanism. Both sexes of the two hummingbird species that inhabit the island are regular visitors. Low fruit and seed set, low genetic diversity, and a shrinking number of populations all contribute to increased concern about the future of this species-and perhaps the hummingbirds that depend on it.     

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.     

Does acoustic priming ‘sweeten the pot’ of floral nectar?

A recent claim that evening primrose flowers adaptively secrete nectar in response to vibrations from hovering bees lacks supporting evidence. The authors fail to demonstrate that bees can access the concealed nectar and that their visits enhance plant fitness. Reanalysis of the authors’ data raises additional concerns about their conclusions.     

Nectar: generation, regulation and ecological functions

Nectar contains water, sugars and amino acids to attract pollinators and defenders and is protected from nectar robbers and microorganisms by secondary compounds and antimicrobial proteins. Floral and extrafloral nectar secretion can be induced by jasmonic acid, it is often adjusted to consumer identity and consumption rate and depends on invertase activity. Invertases are likely to play at least three roles: the uploading of sucrose from the phloem, carbohydrate mobilization during active secretion and the postsecretory adjustment of the sucrose:hexose ratio of nectar. However, it remains to be studied how plants produce and secrete non-carbohydrate components. More research is needed to understand how plants produce nectar, the most important mediator of their interactions with mutualistic animals.     

How common is the ability of extrafloral nectaries to produce nectar droplets,to secrete nectar during the night and to store starch?

Evidence in favour of the ability of extrafloral nectaries (EFNs) to form nectar drop(let)s, secrete extrafloral nectar (EFNec) also during the night and store starch was compiled in order to refute controversial assertions. Not only were more than 150 reports of direct observations of EFNec drop(let)s found, but also 90 studies which suggest that EFNec secretion is copious enough to form drop(let)s automatically by forces of physics (surface tension strength), provided nectar accumulation is not interrupted by predatory animals. Twenty direct observations of nocturnal production of EFNec sufficiently proved that it is not always produced during the day. Additionally, numerous observations of the nocturnal activities of nectar consumers on EFNs indirectly indicated very common nocturnal secretion of EFNec. Although there is an early report of a starch‐containing EFN from 1881 (Trelease), few similar observations in other EFNs followed. Nevertheless, four studies have described the disappearance of stored starch during secretion and senescence of the EFNs. Referring back to an apparent relationship between the degradation of starch stored in a floral nectary and programmed cell death, at least in EFNs with transient storage of starch, a similar relationship cannot be excluded.     

Nectar sugar composition of European Caryophylloideae (Caryophyllaceae) in relation to flower length,pollination biology and phylogeny

Floral nectar composition has been explained as an adaptation to factors that are either directly or indirectly related to pollinator attraction. However, it is often unclear whether the sugar composition is a direct adaptation to pollinator preferences. Firstly, the lower osmolality of sucrose solutions means that they evaporate more rapidly than hexose solutions, which might be one reason why sucrose‐rich nectar is typically found in flowers with long tubes (adapted to long‐tongued pollinators), where it is better protected from evaporation than in open or short‐tubed flowers. Secondly, it can be assumed that temperature‐dependent evaporation is generally lower during the night than during the day so that selection pressure to secrete nectar with high osmolality (i.e. hexose‐rich solutions) is relaxed for night‐active flowers pollinated at night. Thirdly, the breeding system may affect selection pressure on nectar traits; that is, for pollinator‐independent, self‐pollinated plants, a lower selective pressure on nectar traits can be assumed, leading to a higher variability of nectar sugar composition independent of pollinator preferences, nectar accessibility and nectar protection. To analyse the relations between flower tube length, day vs. night pollination and self‐pollination, the nectar sugar composition was investigated in 78 European Caryophylloideae (Caryophyllaceae) with different pollination modes (diurnal, nocturnal, self‐pollination) using high‐performance liquid chromatography (HPLC). All Caryophylleae species (Dianthus and relatives) were found to have nectar with more than 50% sucrose, whereas the sugar composition of Sileneae species (Silene and relatives) ranged from 0% to 98.2%. In the genus Silene, a clear dichotomous distribution of sucrose‐ and hexose‐dominant nectars is evident. We found a positive correlation between the flower tube length and sucrose content in Caryophylloideae, particularly in day‐flowering species, using both conventional analyses and phylogenetically independent contrasts.     

THE POLLINATION ECOLOGY OF AQUILEGIA ELEGANTULA AND A. CAERULEA (RANUNCULACEAE) IN COLORADO

Aquilegia elegantula Greene and A. caerulea James occur in montane and subalpine habitats in the southern Rocky Mountains of western North America. The red and yellow flowers of A. elegantula are nodding, odorless, protogynous, and secrete a concentrated (44%) sucrose nectar in the floral spurs. Seed set in flowers under pollinator exclosures was 12% while seed set in open-pollinated flowers was 65%. The flowers of A. elegantula are pollinated primarily by the Broad-tailed Hummingbird (Selasphorus platycercus [Swainson]) and by at least three species of pollen-foraging bumblebees, of which Bombus occidentalis Greene is the most common. The blue and white flowers of A. caerulea are erect, mildly fragrant, protandrous, and secrete a 26% sucrose nectar. Seed set in caged flowers in the field averaged 39%. in uncaged flowers 54%. The most important pollinators of A. caerulea are the crepuscular hawkmoth, Hyles (=Celerio) lineata (Fabricius) and ten species of pollen-foraging Bombus. The most abundant bumblebee species, B. occidentalis, is also a frequent nectar thief. Differences in pollination systems alone probably do not constitute an effective anti-hydridization mechanism between A. elegantula and A. caerulea, but do serve to reinforce differences in habitat and flowering time that distinguish the two species.     

Induced Floral and Extrafloral Nectar Production Affect Ant‐pollinator Interactions and Plant Fitness

Thousands of plant species throughout tropical and temperate zones secrete extrafloral nectar to attract ants, whose presence provides an indirect defense against herbivores. Extrafloral nectaries are located close to flowers and may modify competition between ants and pollinators. Here, we used Lima bean (Phaseolus lunatus L.) to study the plants interaction between ants and flower visitors and its consequences for plant fitness. To test these objectives, we carried out two field experiments in which we manipulated the presence of ants and nectar production via induction with jasmonic acid (JA). We then measured floral and extrafloral nectar production, the number of patrolling ants and flower visitors as well as specific plant fitness traits. Lima bean plants under JA induction produced more nectar in both extrafloral nectaries and flowers, attracted more ants and produced more flowers and seeds than non‐induced plants. Despite an increase in floral nectar in JA plants, application of this hormone had no significant effects on flower visitor attraction. Finally, ant presence did not result in a decrease in the number of visits, but our results suggest that ants could negatively affect pollination efficiency. In particular, JA‐induced plants without ants produced a greater number of seeds compared with the JA‐treated plants with ants.     

Dual Effect of Phenolic Nectar on Three Floral Visitors of Elsholtzia rugulosa (Lamiaceae) in SW China

Some plants secrete toxic nectar to appeal to most effective pollinators and deter non-pollinators or nectar thieves; however available information about ecological function of toxic nectar remains scarce. Elsholtzia rugulosa stands out as a plant with toxic nectar recorded in SW China. We focused on the functional significance of the phenolic compound that imparts toxic to the nectar of E. rugulosa. The effects of phenolic nectar were studied in three visitors of the flowers of the winter-blooming E. rugulosa Hemsl. (Lamiaceae) in SW China. The pollinating species Apis cerana Fabricius (Apidae; Asian honey bee) and two occasional visitors, Vespa velutina Lepeletier (Vespidae; yellow-legged Asian hornet) and Bombus eximius Smith (Apidae; a bumble bee) were tested for their preferences for low and high concentrations of 4-hydroxybenzoic acid in hexose and sucrose solutions. The pollinator is important for the plant, which is dependent on pollinator visits to attain a higher seed production and it is most likely that the combination of phenolic toxic nectar and the adaptation to phenolic nectar by A. cerana delivers an evolutionary advantage to both actors. The low and high concentrations of the phenolic acid were nearly totally refused by both occasional visitors V. velutina and B. eximius and were preferred by the pollinator A. cerana. E. rugulosa gains by having a much higher seed production and the pollinating honey bee by having an exclusive and reliable food source during the winter season at high altitudes in SW China. We found that the function of the toxic phenolic compound has dual roles by appealing to legitimate pollinators and deterring non-pollinators of E. rugulosa.     

FLORAL BIOLOGY AND BREEDING SYSTEM OF BAUHINIA BENTHAMIANA TAUB. (LEGUMINOSAE), A BAT-POLLINATED TREE IN VENEZUELAN “LLANOS”

The breeding system and floral characteristics of Bauhinia ungulata L. were investigated during two consecutive flowering cycles (Dec-Feb). This tree grows in savannas in small sized patches scattered in the grassland and in larger tree clusters called “mata.” Anthesis occurs between 17:00 and 19:30. Anther dehiscence and nectar secretion begin one hour after anthesis. Nectar is produced for 10 to 12 hr with an average sugar concentration of 13.5%. Flowers in small trees secrete nectar at a higher rate than do flowers in bigger trees. Two phyllostomatid bat species, Glossophaga soricina and Phyllostomus discolor, visit the flowers of B. ungulata between 20:00 and 4:00. Visits of variable duration occur every 1 to 2 hr. In general, the times of greatest nectar production, of highest solute concentration of the nectar and of most frequent visits by the main pollinators coincide. The flowers are significantly dimorphic in pistil length, and ovules are abortive in the short-pistil flowers. Artificial self- and cross-pollinations in long-and short-pistil flowers show that B. ungulata is genetically self-incompatible and functionally andromonoecious.     

Bumblebee visits to Impatiens spp.: pattern and efficiency

Summary Three Japanese species of Impatiens, which secrete nectar continuously in long spurs, were visited by Bombus diversus workers consecutively throughout the day. B. diversus workers showed characteristic patterns of behavior in flower use, flower choice, and patch departure. (1) Bumblebees stayed longer on a flower which had been unvisited for a while than on a flower which had been visited recently. (2) Bumblebees preferred visiting flowers which had been unvisited for a while to visiting those which had been visited recently, and to visiting those which had been unvisited for a long period. (3) Bumblebees had a higher probability of leaving a patch after they had stayed on a flower for a short period than after they had stayed for a longer period. The bumblebees appeared to perceive both remotely and proximately chemical cues deposited by other foraging individuals, which indicated nectar rewards in a flower, and thus obtained a higher nectar intake than the mean amount of nectar left in a flower.     

Bumblebee-pollination and temporal change of the calyx tube length in Clematis stans(Ranunculaceae)

Clematis stans is dioecious semi-arboreal, with pale purple–blue, nodding, tubulous flowers in a paniculate inflorescence. Both male and female flowers produce nectar from the base of the calyx tube during a flowering period of 3 or 4 days, and are pollinated by two bumblebee species, Bombus diversus and B. honshuensis, with different proboscis lengths. When the flowers open, four sepals constructing a calyx tube separate at the top and their respective tips gradually curl up, so that a tubular part shortens. Observations at two field sites showed that B. diversus (with a longer proboscis) most often visits the flowers with a longer calyx tube, and B. honshuensis (with a shorter proboscis) the flowers with a shorter calyx tube, i.e., later in the flowering period. By changing the calyx tube length, the flowers of C. stans accept the two bumblebee species with different proboscis length as pollinators and thus increase the chance of pollination for each flower. It was also found that the two bumblebee species prefer the male flowers to the female flowers, although the female flowers secrete more nectar as a reward than male flowers. This is likely because they visit the male flowers to collect pollen grains in addition to nectar. Electronic Publication     

The timing of nectar secretion in staminal and staminodial glands in Lauraceae

The flowers of many Lauraceae have two kinds of glandular organ: paired glands at the base of the filaments of the third androecial whorl, and staminodes with a glandular head, corresponding to a fourth, sterile androecial whorl. So far, it is unknown why there are two different kinds of organ with apparently the same function. Observations now show that the staminal and the staminodial glands secrete nectar at different times in the heterodichogamous flowering cycle, and are therefore essential for the pollination of bisexual Lauraceae flowers.     

Nectary Structure and Ultrastructure of Unisexual Flowers of Ecballium elaterium(L.) A. Rich. (Cucurbitaceae) and their Presumptive Pollinators

The structure and ultrastructure of the nectaries of the monoeciousspecies Ecballium elaterium were studied. Large differencesin size and structure of the nectaries were observed in thetwo genders of flowers, those of the staminate flowers beingmuch larger and more developed than those of the pistillateflowers. The latter do not secrete measurable amounts of nectar.In the nectariferous cells, especially of the staminate flowers,numerous plasmodesmata are present. The pre-nectar originatingin the phloem is stored in the plastids of the nectariferouscells primarily as starch grains. The nectar appears to be exudedfrom the nectary via modified stomata. Very small insects ofthe order Hemiptera were found to dwell inside the flowers ofthe two genders, but in different numbers; their number in thestaminate flowers was more than twice that in the pistillateflowers. These insects may take part in the process of pollination.Copyright 2001 Annals of Botany Company Ecballium elaterium, Cucurbitaceae, monoecious plant, nectaries, structure, ultrastructure, nectar secretion, stomata, Hemiptera insects     

珍稀濒危植物沙冬青花生物学研究

沙冬青蝶形花冠,柱头具指状突起,分长柱头和短柱头。群落花期约47 d,单花花期约7 d,分为露黄、微开、盛开、凋谢4个时期。TTC法测定4个时期花粉均具活力,可保持70 d左右,苯胺蓝测定柱头可受性3~4 d。沙冬青花一般在开花前一天就开始泌蜜,开花第2~3 d,分泌量达到高峰,花后4~5 d产蜜量减少直至停止,日泌蜜和散粉集中在10:00-14:00。沙冬青开花受环境的影响。     

THE EXTRAFLORAL NECTARIES OF IPOMOEA CARNEA (CONVOLVULACEAE)

Ipomoea carnea (Convolvulaceae) possesses two types of extrafloral nectaries, located on the petiole and on the pedicel. These secrete a complex nectar containing sugars and amino acids. The insects attracted to the extrafloral nectaries are predominantly ants and they are relatively abundant throughout the year. A number of incidents of plant defense as a result of the presence of extrafloral nectary visitors at the extrafloral nectaries of I. carnea were observed and are consistent with the ant-guard theory of the function of extrafloral nectaries.