Analysis of amino acids in nectar from pitchers of Sarracenia purpurea (Sarraceniaceae)

Sarracenia purpurea L. (northern pitcher plant) is an insectivorous plant with extrafloral nectar that attracts insects to a water-filled pitfall trap. We identified and quantified the amino acids in extrafloral nectar produced by pitchers of S. purpurea. Nectar samples were collected from 32 pitchers using a wick-sampling technique. Samples were analyzed for amino acids with reverse-phase high-performance liquid chromatography with phenylisothiocyanate derivatization. Detectable amounts of amino acids were found in each of the 32 nectar samples tested. Mean number of amino acids in a nectar sample was 9 (SD = 2.2). No amino acid was detected in all 32 samples. Mean amount of amino acids in a nectar sample (i.e., amount per wick) was 351.4 ng (SD = 113.2). Nine amino acids occurred in 20 of the 32 samples (aspartic acid, cysteine, glutamic acid, glycine, histidine, hydroxyproline, methionine, serine, valine) averaging 263.4 ng (SD = 94.9), and accounting for ~75% of the total amino acid content. Nectar production may constitute a significant cost of carnivory since the nectar contains amino acids. However, some insects prefer nectar with amino acids and presence of amino acids may increase visitation and capture of insect prey.     

Spectrophotometric analysis of nectar production in Silene vulgaris (Caryophyllaceae)

Significant differences in nectar content were observed between hermaphrodite and female plants of the gynomonoecious-gynodioecious taxon, Silene vulgaris, from populations in northern lower Michigan. These differences in nectar sugar content were observable with a carbohydrate digestion technique involving sulfuric acid, phenol, and measurement of the color change with a spectrophotometer. A hand-held refractometer did not provide sufficient resolution to discern differences between the genders. Both genders were similar in the pattern of sugar content of nectar during floral development. A splitplot factorial design revealed significant differences both among individual plants and stages of floral phenology within each gender. The spectrophotometric technique and statistical analysis provided the needed resolution to discern differences between the genders that we relate to pollinator activity, levels of inbreeding depression, and the maintenance of gynodioecy in this taxon.     

Models of floral pattern in detached flowers of Silene coeli-rosa (L) Godr. (Caryophyllaceae)

Organ number per whorl was analysed in aberrant flowers of the long-day (LD) plant , Silene coeli-rosa , to test a hypothesis that organ number in a whorl takes its cue from an adjacent outer whorl and that perturbed organ number per whorl is not random but defaults to that of closely related taxa or genera of the Caryophyllaceae. When plants were grown under short-days (SD), transferred to LD and the shoot meristem excised and cultured in vitro under SD, the normal pattern of flower development was often disrupted. For example, we observed flowers which comprised floral whorls with an aberrant number of floral organs. In part, this was an effect of tissue culture; however, the over-and-above effect was the establishment of an alternative pattern of development. Our data indicate that two distinct and recurrent patterns occurred in the aberrant flowers we observed in five separate experiments. First, pairs of floral whorls were linked so that aberration in one whorl resulted in the next whorl being more aberrant than normal. Second, the number of organs in aberrant whorls was not random, but defaulted to an organ number which mimicked the flowers of closely related species of Silene or related genera in the Caryophyllaceae.  © 2002 The Linnean Society of London , Botanical Journal of the Linnean Society , 2002, 140 , 229−235.     

Nectar Dynamics and Sugar Composition in Flowers of Silene and Saponaria Species (Caryophyllaceae)

Abstract: Nectar production in Saponaria officincilis and in five species of Silene (S. ciba, S. dioica, S. noctiflora, S. nutans, S. vulgaris ) was examined during two consecutive years (May to July 1993, and May to June 1994) in the Botanical Garden of the University of Giessen. Nectar volume and sugar concentration were studied in relation to time of day, flower sex, flower age, and flowering stage. Nectar amount in all species studied (except S. dioica ) increased in the afternoon or in the evening until midnight (or until the early morning in S. nutans ). After midnight and until midday, nectar volume in non-visited flowers (except S. dioica ) decreased. Nectar volume in non-visited S. dioica flowers increased constantly with flower age, indicating a stable nectar secretion rate, possibly favouring both day- and night-active flower visitors. Even at the time of highest nectar secretion, all species studied presented several nectarless flowers. Sucrose dominance in the nectar of the nocturnal species S. nutans and Saponaria officinalis fits well with the general syndrome of flowers pollinated by hawkmoths. The syndrome also applies to the nocturnal but regularly selfing, S. noctiflora . The more generalis-tic species S. dioica and S. vulgaris , which are regularly visited by bumblebees as well as nocturnal moths, secreted hexose-domi-nant nectar. Unexpectedly, Silene alba , the only nocturnal species that strictly excluded day-active flower visitors by closing flowers during the day, also secreted hexose-dominant nectar. In some cases, nectar volumes and nectar concentration differed significantly between hermaphroditic, male, and female flowers. Female flowers of S. alba, S. dioica , and S. nutans contained significantly less concentrated nectar than male or hermaphroditic ( S. nutans ) ones. In S. noctifiora and S. vulgaris the difference was not statistically significant but nectar concentration did show the same tendency.     

Gender polymorphism in flowers of Silene vulgaris (Moench) Garcke (Caryophyllaceae)

Silene vulgaris occurs in Israel as a stolonifcrous tetraploid which forms mostly sexually polymorphic populations. Fully hermaphrodite and fully male-sterile plants are linked by a range of intermediate forms which are gynomonoecious and bear hermaphrodite, male-sterile, and partially male-sterile flowers. Male-sterile flowers differ from hermaphrodite ones by having a narrower calyx, a shorter corolla, a thicker style, and a considerably larger stigmatic area with longer papillae which are more markedly bulbous basally. In individual anthers of hermaphrodite flowers, diameters of stained pollen grains showed a continuous variation from 36 to 80 μm with a modal value of 56. About 30% of pollen was found to be non-stainable. Reduction in male fertility occurs at different levels, from small or inviable pollen grains through abortion of single anthers, abortion of entire androecia, and through different degrees of gynomonoecy to totally male-sterile individuals. The proportion of stainable pollen to ovules in hermaphrodite flowers alone was found to be less than 100. From the data presented, less than a third of all ovules produced in the different kinds of flower develop into seeds. This may reflect an inadequacy of pollen, which the local population of S. vulgaris can tolerate because of its stoloniferous habit.     

Temporal variation of flower scent in Silene otites (Caryophyllaceae): a species with a mixed pollination system

Floral scent composition and emission are often related to olfactory abilities, preferences and activity times of pollinating agents. However, most studies have focused on species with one pollinator type, and little is known about species with mixed pollination systems. We investigated the temporal variation in flower visitor activity and flower scent emission of Silene otites (Caryophyllaceae), a species regarded as being nocturnal and pollinated by moths and mosquitoes. We found, however, that S. otites is not only visited at night by moths and mosquitoes, but also by a wide range of day‐active insects. Furthermore, both the intensity and composition of floral scent change during the course of the day. Total scent emission reached its maximum in the early hours of the night when compounds known to be attractants for moths and/or mosquitoes, such as lilac aldehyde, benzaldehyde, phenylacetaldehyde and 2‐phenylethanol, dominated the scent. However, although less intensive, emission continued during the day. In the early morning, α‐pinene, hotrienol, 2‐phenylethanol and δ‐3‐carene were the dominant scent compounds. (E)‐β‐Ocimene, hotrienol and benzyl alcohol contributed proportions of at least 10% before midday when flies, such as syrphids, were abundant flower visitors, whereas linalool and hotrienol contributed to the scent with at least 10% each in the afternoon when bees also contributed to flower visits. We conclude that, in S. otites, the changing emission patterns are ‘in tune’ with the olfactory abilities, preferences and activity times of its day‐ and night‐active potentially pollinating flower visitors. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 169 , 447–460.     

The occurrence and significance of amino acids in floral nectar

Approximately 1 500 angiosperm species, in previous papers, have been sampled for the assessment of the amino acids (a. a.) in their nectar. We reaffirm that the findings provide statistically significant data linking differences in the concentration with pollinator type. Flowers that are pollinated by animals that have alternative sources of protein-building a. a.’s show lower a. a. concentration than those that are not. There is a tendency for woody plant nectar a. a.’s to be less concentrated than those of herbaceous plants, but there can be “phylogenetic constraints” which may reduce the correlations of a. a. concentration with pollinator type and with life form. The individual a. a.’s form complements which are qualitatively extremely constant within species. Proline is a normal constituent of many nectars and does not necessarily indicate contamination of the nectar by pollen. Criticism of our findings byGottsberger & al. (1984) is answered by reference to our previous publications and those of other workers, and to the presentation of data from California native species, not published previously. All previous postulates are borne out by these new data with the exception of positive correlations of a. a. concentration with “primitive” and “advanced” floral characteristics taken one at a time, which appear to be inconsistent and are affected strongly by the nature of the family in which they occur. Summary data are provided for families and genera which indicate that high or low a. a. concentration can typify certain families and genera of both relatively “primitive” and relatively “advanced” nature. Needs for future research on an ecosystem basis are quoted.     

Technique for washing nectar from the flowers of Tasmanian leatherwood (Eucryphia lucida Eucryphiaceae)

The technique for extracting floral nectar using micropipettes is often unsuited to flowers where nectar is produced in very small volumes and/or where nectar is highly viscous. An alternative technique for washing the viscous nectar from the flowers of Tasmanian leatherwood, Eucryphia lucida (Labill.) is described. Here, two washes with a known volume of distilled water removed 95% of total floral sugar. Using such a washing technique on exposed and bagged E. lucida flowers provided information on nectar production in this species that could not have been obtained using a standard micropipette extraction method.     

Floral nectar production and nectary anatomy and ultrastructure of Echinacea purpurea (Asteraceae)

BACKGROUND AND AIMS: In spite of the impressive species diversity in the Asteraceae and their widespread appeal to many generalist pollinators, floral-nectary ultrastructure in the family has rarely been investigated. To redress this, a study using Echinacea purpurea, a plant of horticultural and nutraceutical value, was undertaken. Nectar secretion of disc florets was compared with floral nectary ultrastructure taking into account nectar's potential impact upon the reproductive success of this outcrossing species. METHODS: Micropipette collections of nectar in conjunction with refractometry were used to determine the volume and nectar-sugar quantities of disc florets throughout their phenology, from commencement of its production to cessation of secretion. Light, scanning-electron and transmission-electron microscopy were utilized to examine morphology, anatomy and ultrastructure of nectaries of the disc florets. KEY RESULTS: Florets were protandrous with nectar being secreted from anthesis until the third day of the pistillate phase. Nectar production per floret peaked on the first day of stigma receptivity, making the two innermost whorls of open florets most attractive to foraging visitors. Modified stomata were situated along the apical rim of the collar-like nectary, which surrounds the style base and sits on top of the inferior ovary. The floral nectary was supplied by phloem only, and both sieve elements and companion cells were found adjacent to the epidermis; the latter participated in the origin of some of the precursor cells that yielded these specialized cells of phloem. Companion cells possessed wall ingrowths (transfer cells). Lobed nuclei were a key feature of secretory parenchyma cells. CONCLUSIONS: The abundance of mitochondria suggests an eccrine mechanism of secretion, although dictyosomal vesicles may contribute to a granulocrine process. Phloem sap evidently is the main contributor of nectar carbohydrates. From the sieve elements and companion cells, an apoplastic route via intercellular spaces and cell walls, leading to the pores of modified stomata, is available. A symplastic pathway, via plasmodesmata connecting sieve elements to companion, parenchyma and epidermal cells, is also feasible. Uncollected nectar was reabsorbed, and the direct innervation of the nectary by sieve tubes potentially serves a second important route for nectar-sugar reclamation. Microchannels in the outer cuticle may facilitate both secretion and reabsorption.     

The effects of soil fertilizer on amino acids in the floral nectar of corncockle, Agrostemma githago (Caryophyllaceae)

Plants of the meadow annual Agrostemma githago (Caryophyllaceae) were grown in 1-m2 field plots prepared with three fertilizer treatments as follows: (a) "low", no fertilizer, (b) "medium", 75 g of slow release fertilizer granules, (c) "high", 175 g of granular treatment. After sowing in spring the plants were left until flowering in late summer. Nectar was extracted using 5-μl glass microcapillary tubes. The material was frozen and sampled at a later date. The samples were analysed by high performance liquid chromatography (HPLC) using the AccQtag (Waters Corp.) system. Analysis showed that the total concentration of amino acids increased significantly with increasing fertilizer treatment ( P <0.05). Of the amino acids present glutamine showed a large and significant increase ( P <0.01), proline showed an increase ( P <0.05) and γ-amino butyric acid (GABA) showed a decrease ( P <0.05). No others showed a significant change. It is interesting to note that a common biosynthetic pathway (from α-ketoglutarate) links the amino acids that altered in concentration. Due to these changes the relative abundance of about half of the amino acids in the nectar was significantly altered. Glutamine showed a significant increase ( P <0.001) in percentage of the total with most of the remaining amino acids declining in relative abundance. The results show that, in contradiction to earlier work, soil conditions can affect the amino acid complement of nectar. This may have implications for plant-insect interactions, as local populations of pollinators may benefit from the increased amino acid content of the nectar and preferentially visit plants growing in high nutrient conditions.     

Pollination ecology of Silene acutifolia (Caryophyllaceae): floral traits variation and pollinator attraction

BACKGROUND AND AIMS: The floral display influences the composition of pollinators interacting with a plant species. Geographic and temporal variation in pollinator composition complicates the understanding of the evolutionary consequences of floral display variation. This paper analyses the relationships between Silene acutifolia, a hermaphroditic perennial herb, and its pollinators, based on field studies in the north-west of Spain. METHODS: Studies were conducted over three years (1997-1999). Firstly, the main pollinators of this species were determined for two years in one population. Secondly, pollen limitation in fruit and seed production was analysed by supplementary hand pollinations, and counting the pollen grains and tubes growing in styles for two different-sized populations. Finally, the effect of flower size and number on the rate of visitation and total seed number was examined for 15 marked plants. RESULTS AND CONCLUSIONS: The primary pollinators were long-tongued insects, including Hymenoptera, Lepidoptera and Diptera, but the composition and visitation frequencies differed between years. Pollen limitation occurred in one of the years of study. There was between-population variation in the number of pollen grains and pollen tubes found in styles, suggesting pollen limitation in one population. Overall, pollinators visited plants with more open flowers more frequently, and pollinated more flowers within these plants. Conversely, petal and calyx sizes had no effect on insect visitation. Plants with higher rates of visits produced higher number of seeds, suggesting that pollinator-mediated limitation of seed and fruit production may be important in some years.     

Effect of rootstocks on floral nectar composition in apple cultivars

 Insect attraction of flowers in apple cultivars is an essential question from the viewpoint of fruit yield, since apple flowers are autosterile and entomogamous. Therefore, it is important to study the two most significant factors of primary attraction, pollen and nectar production. The composition of nectar in apple cultivars on different rootstocks was investigated, putting an emphasis on sugar components. It was stated that the composition of nectar and the ratio of mono- and disaccharides was almost identical and constant in trees on rootstock M.4, whereas on other rootstocks the effects of the season were more pronounced. In triploid cultivars on rootstocks M.9 and M.26 nectar can be characterised by the dominance of sucrose. Relationship was found between free fertilisation rate of cultivars and sugar amount of nectar. In triploid cultivars nectar sugar content was usually lower than in diploid ones, and reached the bee visitation threshold only in some years, influenced also by the effect of rootstocks. Received August 27, 2002; accepted November 16, 2002 Published online: June 2, 2003     

Breeding system features and a novel method for locating floral nectar secretion in a South American nightshade (Jaltomata quipuscoae)

Abstract

The location of nectar secretion in flowers of Jaltomata has not been identified with certainty until now: removal of the corolla and androecium from one side of living flowers allowed us to see, in progress, nectar secretion by the ovarian nectary. We studied Jaltomata quipuscoae, a wild plant that grows in southern Peru and produces copious, red floral nectar. Unmanipulated flowers do not set fruit in a pollinator-free greenhouse, demonstrating lack of autogamy, but self-compatibility was demonstrated by manual self-pollinations leading to fruit-set. Anther dehiscence is staggered with the anthers of a flower dehiscing over hours on the same day. The corolla and nectar are UV-absorptive. Flowers last 4–10 days, are usually protogynous during the first day the corolla is open, and do not close for the night.     

Nutritional value of floral nectar sources for flight in the parasitoid wasp, Cotesia glomerata

Abstract.  This study investigates the feeding of the parasitoid Cotesia glomerata on the floral nectar of Anethum graveolens and Origanum vulgare and evaluates the nutritional value of these food sources for increasing flight capacity. A significantly higher proportion of females are observed feeding on A. graveolens immediately after being released onto the floral nectar plants compared with those of the group released onto O. vulgare , although female wasps from both groups spent a similar period of time feeding on both nectar sources and there is no significant difference in their body weight increase. Laboratory tests on computer-linked flight mills demonstrate that adult feeding significantly influences flight capacity, as defined by the number of flights, the longest single flight and the total distance flown during a given period of time, in female wasps but not in males, albeit the extent of influence varying with the type of food sources. After ad libitum feeding on the floral nectar of A. graveolens , female wasps perform flight activity as well as those feeding on honey, which is superior to those feeding on sucrose, in all the measured parameters. Feeding on the floral nectar of O. vulgare has a similar effect to feeding on the extrafloral nectar of Vicia faba and significantly increases the total distance flown but not the number of flights or the longest single flight. Feeding on sucrose does not increase flight capacity in the parasitoids beyond those having access to water only. The findings of this study suggest that judicious selection of floral nectar plants and habitat manipulation might play a key role in the biological control of pest insects.     

Variability of nectar production and composition in Linaria vulgaris (L.) Mill. (Scrophulariaceae)

 We studied nectar characteristics during the long flowering period (late June to end of November) in two populations of Linaria vulgaris (L.) Mill. spontaneously growing in the Botanical Gardens of Siena University (Tuscany, central Italy). The two populations were close to each other but they differed in blooming period. Plants of population 1 sprouted in May and flowered from the end of June to the end of September. Population 2 sprouted at the end of August and flowered from September to the end of November. Differences in nectar production and composition were found between and within populations. Flowers of population 1 produced a very small amount of nectar (not collectable) that remained on the nectary surface. The quantity of nectar increased in late September, when each flower produced 2–3 μl of nectar that flowed into the spur. Total sugar concentration was 175.8 mg/ml in young flowers. Flowers of population 2 produced 5–8 μl of nectar with a total sugar concentration of 200.9 mg/ml in the young stage. In bagged senescent flowers nectar volume decreased in both populations and nectar sugar concentration decreased down to 11.6 mg/ml in population 2 and increased up to 289.6 mg/ml in population 1. For both populations, the decrease in nectar volume in bagged flowers may have been due to water loss by evaporation. In population 2, the decrease in sugar concentration may have been due to nectar reabsorption that was never observed in population 1. Nectar variability is discussed in relation to insect visits and seed set. Received August 14, 2002; accepted December 17, 2002 Published online: June 2, 2003     

A new species of Silene (Caryophyllaceae) from east Anatolia, Turkey

Silene bitlisensis O.Tugay & Ertugrul from the Green Line Afforestation Zone (B9: Bitlis Province) is a new species described and illustrated here. It is closely related to S. caramanica Boiss. & Heldr. var. ilarslanii Aytaç & Dural, from which it mainly differs in its habit, leaf, and floral features.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 463–466.