An Effect of Ethylene on the Distribution of 14C-sucrose from the Petals to Other Flower Parts in the Senescent Cut Inflorescence of Dianthus caryophyllus

The distribution of carbon-14 in the flower parts of the cutcarnation inflorescence after feeding ¹⁴C-sucrose through thepetals was studied during natural ageing and after ethylenetreatment. Levels of ethylene which caused irreversible wiltingof petals also promoted an accelerated transfer of the radioactivesucrose to the nectar, gynaecium and stem. Since the nectarreceived a relatively large proportion of the radioactive carbon,the composition of the sugars in the nectar and the vascularizationof the nectary were investigated. Sucrose comprised about 85per cent of the nectar sugars and the balance was glucose andfructose. The vascular tissue closest to the nectary consistedof phloem elements; tracheary elements terminated deeper inthe receptacle and were surrounded by a ring of phloem. Thepercentage of solutes in the nectar was about 18 per cent andincreased when the flowering stems were placed in sucrose solutions;the solutes in the nectar were principally sugars. Taken togetherthe results show that the nectary can act as a sink for sucroseand, in the flower at least, that translocation of sucrose takesplace in the phloem. The results provide further evidence forthe hypothesis that ethylene promotes mobilization of substrateand an efflux of material from petals to the gynaecium, nectarand stem.     

Malonylated flavonol glycosides from the petals of Clitoria ternatea

Three flavonol glycosides, kaempferol 3-O-(2"-O-alpha-rhamnosyl-6"-O-malonyl)-beta-glucoside, quercetin 3-O-(2"-O-alpha-rhamnosyl-6"-O-malonyl)-beta-glucoside, and myricetin 3-O-(2",6"-di-O-alpha-rhamnosyl)-beta-glucoside were isolated from the petals of Clitoria ternatea cv. Double Blue, together with eleven known flavonol glycosides. Their structures were identified using UV, MS, and NMR spectroscopy. They were characterized as kaempferol and quercetin 3-(2(G)- rhamnosylrutinoside)s, kaempferol, quercetin, and myricetin 3-neohesperidosides, 3-rutinosides, and 3-glucosides in the same tissue. In addition, the presence of myricetin 3-O-(2"-O-alpha-rhamnosyl-6"-O-malonyl)-beta-glucoside was inferred from LC/MS/MS data for crude petal extracts. The flavonol compounds identified in the petals of C. ternatea differed from those reported in previous studies.     

Spatial fragrance patterns in flowers of Silene latifolia: Lilac compounds as olfactory nectar guides?

Floral odour can differ qualitatively and quantitatively between different parts of the flowers, and these spatial fragrance patterns within the flowers can be used by pollinators for orientation on flowers. Here we present results of spatial fragrance patterns within flowers of the dioecious Silene latifolia (Caryophyllaceae). Volatiles were collected and analysed using a highly sensitive dynamic headspace method, which allows dramatically reducing the sample time. From all flower parts, especially the petals and the anthophore emitted the typical flower volatiles of S. latifolia. However, compounds emitted from the petals differed from compounds emitted by the anthophore. The anthophore emitted the monoterpenoids lilac aldehydes and alcohols, whereas, all other typical scent compounds (e.g. benzoids, other monoterpenoids) were emitted by the petals. Lilac aldehydes are known to be behaviourally very attractive for noctuid moths, and they may serve as nectar guides in S. latifolia.     

Bulk Segregant Analysis of an Induced Floral Mutant Identifies a MIXTA-Like R2R3 MYB Controlling Nectar Guide Formation in Mimulus lewisii

The genetic and developmental basis of many ecologically important floral traits (e.g., carotenoid pigmentation, corolla tube structure, nectar volume, pistil and stamen length) remains poorly understood. Here we analyze a chemically induced floral mutant of Mimulus lewisii through bulk segregant analysis and transgenic experiments and identify a MIXTA-like R2R3 MYB gene that controls nectar guide formation in M. lewisii flowers, which involves epidermal cell development and carotenoid pigmentation.     

Floral biology and pollination in stapeliads — new results and a literature review

The myiophilous pollination syndrome of stapeliads is investigated. Olfactory stimuli, imitating dung or decaying organic (zoogenic or phytogenic) matter, are used as attractants, together with mimetic colouration and, sometimes mimetic sculpturing. In most species nectar is present. The nectar mainly serves as optical attractant causing brilliance effects, and as visitor guide. However, nectar obviously is also a reward in many species. Flies (Cyclorrhapha:Calyptratae) constitute the most important pollinating group. In the pollination process they carry pollinaria only at the distal parts of their proboscis, never on their legs. No fundamental differences between the pollinator spectra of flowers in habitat and cultivated ones were observed.     

Formation of UV-honey guides in Rudbeckia hirta

The UV-honey guides of Rudbeckia hirta were investigated by UV-photography, reflectance spectroscopy, LC-MS analysis and studies of the enzymes involved in the formation of the UV-absorbing flavonols present in the petals. It was shown for the first time that the typical bull’s eye pattern is already established at the early stages of flower anthesis on the front side of the petal surface, but is hidden to pollinators until the buds are open and the petals are unfolded. The rear side of the petals remains UV-reflecting during the whole flower anthesis. Studies on the local distribution of 19 flavonols across the petals confirmed that the majority are concentrated in the basal part of the ray flower. However, in contrast to the earlier studies, eupatolitin 3-O-glucoside (6,7-dimethoxyquercetin 3-O-glucoside) was present in both the basal and apical parts of the petals, whereas eupatolin (6,7-dimethoxyquercetin 3-O-rhamnoside) was exclusively found in the apical parts. The enzymes involved in the formation of the flavonols in R. hirta were demonstrated for the first time. These include a rare flavonol 6-hydroxylase, which was identified as cytochrome P450-dependent monooxygenase and did not accept any methylated flavonol as substrate. All enzymes were present in the basal and apical parts of the petals, although some of them clearly showed higher activities in the basal part. This indicates that the local accumulation of flavonols in R. hirta is not achieved by a locally restricted presence of the enzymes involved in flavonol formation.     

Spatial and temporal patterns of floral scent emission in Dianthus inoxianus and electroantennographic responses of its hawkmoth pollinator

Scent emission is important in nocturnal pollination systems, and plant species pollinated by nocturnal insects often present characteristic odor compositions and temporal patterns of emission. We investigated the temporal (day/night; flower lifetime) and spatial (different flower parts, nectar) pattern of flower scent emission in nocturnally pollinated Dianthusinoxianus, and determined which compounds elicit physiological responses on the antennae of the sphingid pollinator Hyles livornica.The scent of D.inoxianus comprises 68 volatile compounds, but is dominated by aliphatic 2-ketones and sesquiterpenoids, which altogether make up 82% of collected volatiles. Several major and minor compounds elicit electrophysiological responses in the antennae of H. livornica. Total odor emission does not vary along day and night hours, and neither does along the life of the flower. However, the proportion of compounds eliciting physiological responses varies between day and night. All flower parts as well as nectar release volatiles. The scent of isolated flower parts is dominated by fatty acid derivatives, whereas nectar is dominated by benzenoids. Dissection (= damage) of flowers induced a ca. 20-fold increase in the rate of emission of EAD-active volatiles, especially aliphatic 2-ketones.We suggest that aliphatic 2-ketones might contribute to pollinator attraction in D. inoxianus, even though they have been attributed an insect repellent function in other plant species. We also hypothesize that the benzenoids in nectar may act as an honest signal (‘nectar guide’) for pollinators.     

Insect Attracting Structures on Erodium Petals (Geraniaceae)

Abstract: Two types of structures previously unrecorded in Erodium petals are investigated. Spherical hairs filled with liquid resembling nectar droplets are exclusive to an ibero-mauritanic group of species included in Erodium subsect. Romana. Broad flat hairs which reflect light, shining as do nectar droplets, are restricted to most of the species included in Erodium sect. Malacoidea. Long, simple hairs in petals and sepals are involved in collection of nectar droplets. Some of them are arranged at the margin of the petal claw, just over the nectaries. Others are on the internal surface of sepals or on the upper surface of petals, serving apparently the same function. Their shape is aciculate or flattened. The nectar collected among the hairs forms shining spherical droplets, perceptible to insects. The glistening flat hairs and spheres shine in a similar way, probably mimicking nectar and attracting insects. Species with these special nectar-like structures produce nectar in quantities that can be observed by the naked eye, suggesting that these structures increase the attraction efficiency of flowers. Some taxonomic and biogeographic consequences are also discussed.     

Effect of zinc on nectar secretion of Hibiscus rosa-sinensis L

Zinc toxicity in secretory cells caused a range of effects, mainly depending on metal concentration. Low concentrations activated nectary function increasing nectar secretion but secretion was greatly inhibited or stopped entirely by ongoing concentration. Water loss rate of zinc treated flower parts was significantly reduced whereas green sepals were dehydrated more rapidly in comparison to colored petals. The content of zinc, calcium, magnesium and manganese increased mainly in sepals under excess of zinc, but in the secreted nectar this metal was not evident. Morphological changes were observed in mucilage cells concerning the mucilage structure and appearance. The parenchymatic, subglandular cells displayed an early vacuolarization and cytoplasm condensation. Secretory hairs appeared to be thinner, the apical cell folded inwards and plasmolytic shrinkage became severe in all cells. The waxy cuticula showed an increased electron density. A plasmalemma detachment from the external cell walls was observed creating a gap between cell wall and plasmalemma. ER cisterns of all treated nectary hairs dominated the cytoplasm and electron dense deposits were seen within its profiles. A great number of other organelles were also present, showing electron dense deposits in their membranes as well. The vacuome was drastically reduced in all cells, except in the subglandular ones and electron dense membrane remnants were observed.     

Predation by blue crabs, Callinectes sapidus, on rapa whelks, Rapana venosa: possible natural controls for an invasive species?

Blue crabs Callinectes sapidus are voracious predators in Chesapeake Bay and other estuarine habitats. The rapa whelk Rapana venosa is native to Asian waters but was discovered in Chesapeake Bay in 1998. This predatory gastropod grows to large terminal sizes (in excess of 150 mm shell length (SL)) and has a thick shell that may contribute to an ontogenetic predation refuge. However, juvenile rapa whelks in Chesapeake Bay may be vulnerable to predation by the blue crab given probable habitat overlap, relative lack of whelk shell architectural defenses, and the relatively large size of potential crab predators. Feeding experiments using three size classes of blue crab predators in relation to a size range of rapa whelks of two different ages (Age 1 and Age 2) were conducted. Blue crabs of all sizes tested consumed Age 1 rapa whelks; 58% of all Age 1 whelks offered were eaten. Age 2 rapa whelks were consumed by medium (67% of whelks offered were eaten) and large (70% of whelks offered were eaten) blue crabs but not by small crabs. The attack methods of medium and large crabs changed with whelk age and related shell weight. Age 1 whelks were typically crushed by blue crabs while Age 2 whelk shells were chipped or left intact by predators removing prey. Rapa whelks less than approximately 35 mm SL are vulnerable to predation by all sizes of blue crabs tested. Rapa whelk critical size may be greater than 55 mm SL in the presence of large blue crabs indicating that a size refugia from crab predation may not be achieved by rapa whelks in Chesapeake Bay until at least Age 2 or Age 3. Predation by blue crabs on young rapa whelks may offer a natural control strategy for rapa whelks in Chesapeake Bay and other estuarine habitats along the North American Atlantic coast.     

On the function of pea flower feeding by Bruchus pisorum

Pea flower feeding by adult pea weevils, Bruchus pisorum (L.) (Coleoptera: Bruchidae), with special emphasis on nectar feeding, was investigated in a series of laboratory experiments. Male and female adults robbed nectar from flowers of the garden and field pea, Pisum sativum L., and females which fed on the nectar, petals, and female organs of pea flowers lived significantly longer than those denied food and water and those that fed on water only. The results of other experiments suggested that pea flower qualities other than pollen influenced the reproductive success of female B. pisorum. It is hypothesized that pollen seeking B. pisorum effected cross-pollination in the wild progenitor of the modern-day autogamous pea, and adult pea weevils of both sexes rob pea nectar to obtain a readily available source of energy to sustain flight.     

Nectar production and carbohydrate composition across floral sexual phases: contrasting patterns in two protandrous Aconitum species (Delphinieae,Ranunculaceae)

In dichogamous species, it is necessary that pollinators are attracted both to male- and female-phase flowers in order to achieve effective cross-pollination. We investigated, over a two-year period, how nectar production and composition differs in protandrous flowers of Aconitum lycoctonum L. and Aconitum carmichaelii Debeaux, two species originating from different geographical regions. Flowers of A. carmichaelii secreted approx. 2.5-fold more nectar than flowers of A. lycoctonum. The nectar of A. carmichaelii was sucrose-dominant (87.6:9.5:2.9, sucrose:fructose:glucose), whereas sucrose-rich nectar, lacking glucose, occurred in A. lycoctonum (39.9:60.1, sucrose:fructose). Total sugar concentration was similar (53%, on average) in both species. These species also showed contrasting patterns in nectar production and composition between the floral sexual phases. On average, in A. carmichaelii, male-phase flowers produced 2.4-fold more nectar than female-phase flowers in the two years of study, and nectar was sucrose-dominant, irrespective of sexual phase. By contrast, nectar production in A. lycoctonum was biased towards the male phase in one year of study and towards the female phase in the other, and whereas nectar was sucrose-dominant during the male-phase, it was fructose-rich during the female phase, indicating sucrose re-absorption. Although the characteristics of nectar in these Aconitum species indicate pollination by bumblebees, it is possible that their biogeographical history, and not pollinator selection alone, is important in understanding the lack of glucose in A. lycoctonum nectar. Variability in nectar production and/or carbohydrate composition between sexual phases suggests indirectly that nectar traits complement each of the latter. Nevertheless, further research is required if we are to understand the significance of these disparities in pollination ecology, i.e. for promoting pollinator movements between flowers and plants in order to achieve cross-pollination and avoid inbreeding.     

Expression of floral MADS-box genes in Sinofranchetia chinensis (Lardizabalaceae): implications for the nature of the nectar leaves

Background and Aims

The perianths of the Lardizabalaceae are diverse. The second-whorl floral organs of Sinofranchetia chinensis (Lardizabalaceae) are nectar leaves. The aim of this study was to explore the nature of this type of floral organ, and to determine its relationship to nectar leaves in other Ranunculales species, and to other floral organs in Sinofranchetia chinensis.

Methods

Approaches of evolutionary developmental biology were used, including 3′ RACE (rapid amplification of cDNA ends) for isolating floral MADS-box genes, phylogenetic analysis for reconstructing gene evolutionary history, in situ hybridization and tissue-specific RT-PCR for identifying gene expression patterns and SEM (scanning electron microscopy) for observing the epidermal cell morphology of floral organs.

Key Results

Fourteen new floral MADS-box genes were isolated from Sinofranchetia chinensis and from two other species of Lardizabalaceae, Holboellia grandiflora and Decaisnea insignis. The phylogenetic analysis of AP3-like genes in Ranunculales showed that three AP3 paralogues from Sinofranchetia chinensis belong to the AP3-I, -II and -III lineages. In situ hybridization results showed that SIchAP3-3 is significantly expressed only in nectar leaves at the late stages of floral development, and SIchAG, a C-class MADS-box gene, is expressed not only in stamens and carpels, but also in nectar leaves. SEM observation revealed that the adaxial surface of nectar leaves is covered with conical epidermal cells, a hallmark of petaloidy.

Conclusions

The gene expression data imply that the nectar leaves in S. chinensis might share a similar genetic regulatory code with other nectar leaves in Ranunculales species. Based on gene expression and morphological evidence, it is considered that the nectar leaves in S. chinensis could be referred to as petals. Furthermore, the study supports the hypothesis that the nectar leaves in some Ranunculales species might be derived from stamens.     

Flower morphology,nectar traits and pollinators of Cerinthe major (Boraginaceae-Lithospermeae)

In April and May 2010, a natural population of Cerinthe major (Boraginaceae-Lithospermeae) was investigated with regard to: floral morphology, phenology, sexual receptivity during anthesis, the production of nectar and its sugar and amino acid content. The pollinator array of this species was also investigated. Cerinthe major has showy, pentamerous, hermaphroditic flowers with a deep yellow corolla tube arranged on numerous scorpioid inflorescences. Nectar and pollen are the main floral rewards for pollinators. The arrangement of stamens limits access to nectar and promotes pollen loading onto pollinator bodies as they force their way towards the source of nectar. By limiting the direct exposure of nectar to the environment, the corolla tube and the arrangement of anther filaments seemingly protect nectar from evaporation resulting from high temperatures and low relative humidity during flower anthesis. The green, annular nectary located at the base of the ovary lobes is composed of a very thin epidermis enclosing a dense mass of parenchyma. The epidermis lacks stomata, and the thin cuticle has an irregular surface. The parenchyma cells do not store starch during the pre-secretory stage, suggesting that immediate photosynthesis is the most likely source of nectar carbohydrates. Generally, anthesis lasts 4–5 days; the gynoecium becomes fully receptive on the second day and this is synchronous with anther dehiscence. Since nectar production begins during the bud stage, a substantial volume is available for flower visitors at anthesis. Nectar production decreases drastically in senescent flowers and, if not consumed, the nectar can be re-absorbed. Analysis of sugars revealed a sucrose-dominant nectar (sucrose = 93.18 ± 1.35%). Proline, GABA, taurine, leucine, citrulline and alanine were the main amino acids present and are probably implicated in pollinator flight performance.     

Cytokinins in cut carnation flowers. II. Relationship between endogenous ethylene and cytokinin levels in the petals

Tentative identification using HPLC and RIA techniques indicated the presence of zeatin-O-glucoside, zeatin, ribosylzeatin, dihydrozeatin, iso-pentenyladenine and iso-pentenyladenosine in the petals of carnation flowers. Dihydrozeatin is apparently responsible for most of the biological activity. Within the petals most activity was detected in the basal parts which also senesced much slower than the upper parts of the petals. Treatment with AOA extended petal longevity and reduced ethylene production. This was associated with higher cytokinin-like activity in the basal parts of the petals.These higher levels of cytokinins were not observed in the petals of ACC treated flowers or in the detached control flowers. It is suggested that cytokinin transport and/or metabolism may play an important role in regulating ethylene production in cut carnations.     

Amino acids and protein profile in floral nectar: Much more than a simple reward

Nectar is considered a primary alimentary reward for a large variety of pollinators. Recent studies demonstrated that nectar may have other functions in addition to attracting pollinators. Mainly other two functions have been recognized: defense against microbial invasion and post-secretion modification of sugar profile. The floral nectar of Cucurbita pepo L. has been analyzed to confirm the presence of defense proteins, namely xylosidases that were identified by means of a proteomic approach in a previous study. An enzyme assay was also performed that revealed low invertase activity for which the optimal pH was determined. This invertase activity is not able to modify the sugar profile significantly during the short period of anthesis (6 h). The amino acids complement of the nectar of both sexes was also determined. Sixteen of the 20 protein amino acids have been detected. Proline comprises more than 30% of total amino acid content in male and female flowers. Three non-protein amino acids (taurine, β-alanine, and GABA) represent almost one third of the total amino acid content, and of these, GABA is the most abundant (16%). Several biological activities were attributed to these amino acids and further studies are needed to assess their presumed involvement in managing the foraging behavior of insects. More recent studies, including our own, demonstrate that the various functions of nectar are much more complex than previously thought.     

Changes in 1-aminocyclopropane-1-carboxylic-acid content of cut carnation flowers in relation to their senescence

The rise in ethylene production accompanying the respiration climacteric and senescence of cut carnation flowers (Dianthus caryophyllus L. cv. White Sim) was associated with a 30-fold increase in the concentration of 1-aminocyclopropane-1-carboxylic acid (ACC) in the petals (initial content 0.3 nmol/g fresh weight). Pretreatment of the flowers with silver thiosulfate (STS) retarded flower senescence and prevented the increase in ACC concentration in the petals. An increase in ACC in the remaining flower parts, which appeared to precede the increase in the petals, was only partially prevented by the STS pretreatment. Addition of aminoxyacetic acid (2 mM) to the solution in which the flowers were kept completely inhibited accumulation of ACC in all flower parts.Abbreviations ACC 1-aminocyclopropane-1-carboxylic acid - AOA -aminoxyacetic acid - STS silver thiosulfate complex     

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.     

扫帚菜-白菜轮作对白菜镉吸收的影响

通过盆栽和大田试验研究扫帚菜Kochia scoparia对土壤中镉(Cd)的富集效率,并利用盆栽试验将其与4个白菜品种进行轮作试验,验证扫帚菜对土壤Cd污染的修复效果。结果表明,扫帚菜各部位富集能力表现为叶>根>茎,富集系数分别为15.07、5.44和2.96;种植扫帚菜后土壤总Cd降低6.02%-13.60%;土壤脲酶和酸性磷酸酶活性也有所提高。盆栽轮作结果表明,扫帚菜-白菜轮作系统中白菜地上部Cd含量与未轮作的对照组相比平均降低17.21%,生物量有略微增加,地上部对Cd的转运系数无明显变化。结果显示,通过扫帚菜与白菜轮作不仅可以增加白菜产量,而且可以有效降低白菜可食部Cd含量,实现边生产边治理的绿色农业理念。