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Nectary of Anagyris foetida . A , Cross section of the middle part of the hypanthium stained with periodic acid–Schiff reagent (PAS) and amido black showing the vascular bundles that irrigate the nectariferous parenchyma and the stamens, the vascular bundles that irrigate the calyx and corolla, and the nectariferous parenchyma. B , Cross section stained with PAS showing the nectariferous parenchyma and two vascular bundles. C , View under fluorescence microscopy of the nectary cuticle stained with auramine O. D , Longitudinal section of the hypanthium stained with toluidine blue and ruthenium red showing the extent of the nectariferous tissue. Inset , nectary stomata. Ca 1⁄4 calyx; Co 1⁄4 corolla; Cp 1⁄4 chlorophyllic parenchyma; Cu 1⁄4 cuticle; Ep 1⁄4 epidermis; Nc 1⁄4 nectar cavity; S 1⁄4 stipe; St 1⁄4 stamen; Vb 1⁄4 calyx and corolla vascular bundles; Vs 1⁄4 nectary and stamen vascular bundles. Asterisk indicates stomata pore. Scale bars: 500 m m ( A , D ), 150 m m ( B ), 50 m m ( C ).
Source publication
In Anagyris foetida, a shrubby legume with autumn–winter flowering, the flowers produce great amounts of very dilute nectar during the first half of their life, consonant with their pollination by passeriforms. With advancing age, the volume of nectar diminishes and the concentration increases to values characteristic of bee-pollinated flowers. The...
Contexts in source publication
Context 1
... foetida flowers are papilionaceous, pendulous, odorless, greenish yellow when young, and yellow when old. The standard is smaller than the wings and the keel. The an- droecium is formed by 10 free stamens, and the gynoecium, with seven to 10 ovules, is located on a stipe at the base of a small hypanthium, where the nectar is accumulated (see fig. 1A, 1B in Ortega- Olivencia et al. 2005). The flowers last 10- 14 d if they are not visited (less when pollinated), with this longevity also related to the environmental conditions (F. J. Valtueñ a, A. Ortega-Olivencia, and T. Rodríguez-Riañ o, un- published ...
Context 2
... study of the nectary. The flower nectary con- sists of a disk that covers the inner surface of the hypanthium from its lower part to the base of the insertion of the stamens ( fig. 1). This is therefore a nectary that is not exposed to the light and of a deep green color. The cross sections of the nec- tary showed a single-layer epidermis externally covered by a continuous, thick cuticle in which no pores were observed ( VALTUEN˜AVALTUEN˜VALTUEN˜A ET AL.-NECTAR PRODUCTION IN ANAGYRIS FOETIDA zone of the hypanthium ...
Context 3
... ( fig. 1). This is therefore a nectary that is not exposed to the light and of a deep green color. The cross sections of the nec- tary showed a single-layer epidermis externally covered by a continuous, thick cuticle in which no pores were observed ( VALTUEN˜AVALTUEN˜VALTUEN˜A ET AL.-NECTAR PRODUCTION IN ANAGYRIS FOETIDA zone of the hypanthium ( fig. 1D, inset). They were closed in preanthesic flowers and open during the secretory phase and in postanthesic ...
Context 4
... production in this species was characterized by an initial phase of secretion followed by another of cessation, af- ter which the volume diminished until the flower withered ( fig. 4A1), as did the sugar content ( fig. 4B1). Both parame- ters peaked in the first half of anthesis-between days 1 and 3 (or 4, depending on the year)-independent of whether the flower was visited once or thrice daily ( fig. 4A1, 4B1; data for the 2003-2004 period are similar and are not shown). A repeated-measures nested ANOVA showed that ...
Context 5
... production in this species was characterized by an initial phase of secretion followed by another of cessation, af- ter which the volume diminished until the flower withered ( fig. 4A1), as did the sugar content ( fig. 4B1). Both parame- ters peaked in the first half of anthesis-between days 1 and 3 (or 4, depending on the year)-independent of whether the flower was visited once or thrice daily ( fig. 4A1, 4B1; data for the 2003-2004 period are similar and are not shown). A repeated-measures nested ANOVA showed that both the nec- tar volume and the ...
Context 6
... initial phase of secretion followed by another of cessation, af- ter which the volume diminished until the flower withered ( fig. 4A1), as did the sugar content ( fig. 4B1). Both parame- ters peaked in the first half of anthesis-between days 1 and 3 (or 4, depending on the year)-independent of whether the flower was visited once or thrice daily ( fig. 4A1, 4B1; data for the 2003-2004 period are similar and are not shown). A repeated-measures nested ANOVA showed that both the nec- tar volume and the amount of sugar produced daily differed between days (volume: F ¼ 52:79, P < 0:001; sugar: F ¼ 22:58, P < 0:001) and between years (volume: F ¼ 7:58, P < 0:001; sugar: F ¼ 8:01, P < 0:001). ...
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In this study we characterized the volume, concentration and sugar composition of floral nectar in Anarthrophyllum desideratum (DC) Benth., an endemic shrub from the Patagonian steppe pollinated by Passerine birds, and we evaluated it spatio-temporal variation. We analyzed the differences among ecological areas, populations, plants and years, and t...
Differences in nectar characteristics of bird-pollinated plants were originally attributed to
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Subsequently this has been refuted and it is now hypothesized that these differences reflect
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Summary: Characterization and spatio-temporal variation of nectar in Anarthrophyllum desideratum (Fabaceae): Influence of climate and pollinators. In this study we characterized the volume, concentration and sugar composition of floral nectar in Anarthrophyllum desideratum (DC) Benth., an endemic shrub from the Patagonian steppe pollinated by Passe...
Differences in nectar characteristics of bird-pollinated plants were originally attributed to whether the plant was pollinated by non-passerine hummingbirds or passerine nectarivores. Subsequently this has been refuted and it is now hypothesized that these differences reflect whether the plants are pollinated by occasional or specialist nectarivore...
Citations
... It has a foul smell and yellowish-green color in its young stage but is green in its mature stage (Ortega-Olivencia et al., 2005). Its flowers are important for beekeeping due to their secretions (Valtueña, Ortega-Olivencia & Rodríguez-Riaño, 2007). It is typically found in the coastal areas of Türkiye. A. foetida is avoided by ruminant animals because of the odor produced by the cytisine alkaloid and goats are the only animal known to graze on it. ...
Background
This study evaluated the extent to which the endemic herbaceous and woody species of shrubby rangelands met the roughage needs of grazing animals throughout the year.
Methods
The biomass, botanical composition, and quality of hay were investigated in the shrubby rangelands in Paşaköy of the Ayvacık districts in Çanakkale over the course of a year. Plant samples were taken from the herbaceous species monthly and from the grazing parts of the shrubs in May and November.
Results
The total amount of biomass (hay + shrub) in the rangeland was found to be 30.448 kg/ha. Shrubs made up 18.78% of the rangeland, while the annual species comprised 54.96%, and perennial herbs covered 26.26% of the total biomass. Crude protein (CP) ratios of herbaceous species decreased continuously from March (13.58%) to September (6.73%), and then increased. A similar change was also seen in pure ash (PA) ratios. The CP ratios in the shrub species were high in spring and decreased in autumn and there was an irregular variation in PA rates. Oak had the highest PA ratio during the spring, while thuja had the highest ratio in autumn, and Juniperus oxycedrus during the winter months. In herbaceous species, cell wall components (NDF, ADF, and ADL) reached their highest levels in summer and decreased in spring and winter. However, in shrubs, these components varied according to the species and were generally lowest in spring and then increased in autumn and winter. Here, it was determined that year-round grazing is a suitable grazing system in the shrubby rangelands of the Mediterranean zone, and animals are able to find fresh forage in the rangelands due to the presence of shrubs. However, since the contribution of shrubs to the total forage production is low, additional roughage should be provided, except in the spring when the production and quality of hay increase. These practices may contribute to better livestock management.
... etion of Satsuma mandarin due to the particular structure of the flower. The largest amount of nectar was formed in 2014, when the air temperature was between 14.7 ºC and 16.9 ºC and the average relative humidity was above 80%. In addition to meteorological conditions for nectar secretion, other factors such as flower age (Torres and Galetto, 1998;Valtuena, et. al., 2007), flower size (Longo and Fischer, 2006), plant density (Klinkhamer, 2004), flower sex (Liu et al., 2002;Carlson, 2007), time of day (Macukanovic-Jocic, et.al, 2004;Valtuena et al., 2007), and number of pollinators (Torres and Galetto, 1998;Valtuena, et al., 2007;Keasar et. al., 2008). ...
... In addition to meteorological conditions for nectar secretion, other factors such as flower age (Torres and Galetto, 1998;Valtuena, et. al., 2007), flower size (Longo and Fischer, 2006), plant density (Klinkhamer, 2004), flower sex (Liu et al., 2002;Carlson, 2007), time of day (Macukanovic-Jocic, et.al, 2004;Valtuena et al., 2007), and number of pollinators (Torres and Galetto, 1998;Valtuena, et al., 2007;Keasar et. al., 2008). ...
... In addition to meteorological conditions for nectar secretion, other factors such as flower age (Torres and Galetto, 1998;Valtuena, et. al., 2007), flower size (Longo and Fischer, 2006), plant density (Klinkhamer, 2004), flower sex (Liu et al., 2002;Carlson, 2007), time of day (Macukanovic-Jocic, et.al, 2004;Valtuena et al., 2007), and number of pollinators (Torres and Galetto, 1998;Valtuena, et al., 2007;Keasar et. al., 2008). ...
... Similarly, chloroplasts with a typical structure and well-developed thylakoids were observed in green nectaries of other plants (Zer and Fahn 1992;Fahn and Simony 2001;Vesprini et al. 2012). Additionally, as suggested by Valtuena et al. (2007) and Lüttge (2013), the secretion rates in photosynthetic nectaries seem to be high and sufficient even for pollinators that require substantial amounts of nectar. These authors claim that the photosynthesis in nectariferous tissue is a beneficial phenomenon, as it results in not only production of carbohydrates, which may greatly increase the nectar yield, but also production of energy required for secretion of nectar. ...
... These authors claim that the photosynthesis in nectariferous tissue is a beneficial phenomenon, as it results in not only production of carbohydrates, which may greatly increase the nectar yield, but also production of energy required for secretion of nectar. Similar to R. viscosa, photosynthetic nectaries are present in the flowers of other representatives of Papilionoideae, e.g. in Vicia faba (Davis and Gunning 1992;Stpiczyńska 1995), Pisum sativum (Razem and Davis 1999), Glycine max (Horner et al. 2003) and Anagyris foetida (Valtuena et al. 2007). ...
... Scale bars = 1 μm (a, b, e, f), 500 nm (c, d) transferred from other parts of the flower via phloem strands, which are branches of the hadrocentric vascular bundles (stamen traces) located in close proximity to glandular parenchyma. Previous studies of other species from the Fabaceae family have shown uptake of carbohydrates from phloem sap leaking from adjacent filament bundles, corolla petals, sepals, flower pedicels, receptacle tissues and adjacent leaves (Davis et al. 1988;Razem and Davis 1999;Valtuena et al. 2007;Paiva and Machado 2008). Many researchers indicate that phloem photosynthates are the primary (predominant) source of pre-nectar carbohydrates necessary for nectar production. ...
Floral nectaries are important components of floral architecture and significant taxonomic traits facilitating assessment of relationships between taxa and can contribute substantially to studies on the ecology and evolution of a particular genus. Knowledge of nectary structure and functioning allows better understanding of the mutualistic interactions between the pollinator and the plant. Robinia viscosa var. hartwigii (Hartweg’s locust), planted in many European countries as an ornamental plant and used for recovery of degraded areas and urban arborisation, is a valuable melliferous species often visited by honeybees and bumblebees. The aim of this study was to investigate the microstructure of the floral nectaries of R. viscosa var. hartwigii with the use of light, fluorescence, scanning, and transmission electron microscopes. The photosynthetic nectaries were located on the inner surface of the cup-like receptacle. The components of pre-nectar were synthesised in the chloroplasts of the glandular parenchyma and transported via the conducting elements of the phloem. Nectar was released through modified nectarostomata. Nectar secretion presumably proceeded in the eccrine mode, whereas nectar transport represented the symplastic and apoplastic types. The cuticle on the nectary epidermis surface contained lipids, essentials oils, and flavonoids, while proteins and flavonoids were present in the glandular parenchyma cells. Idioblasts containing phenolic compounds, tannins, and polysaccharides were observed between the glandular parenchyma cells. The location of the nectaries and the mode of nectar production in the flowers of the Hartweg’s locust follow the common location and structure pattern characteristic for the nectaries in some members of the subfamily Papilionoideae and can be a significant taxonomic trait for the genus Robinia and the tribe Robinieae.
... Anagyris foetida is a leguminous shrub or small tree, deciduous in summer, with a Mediterranean and Irano-Turanian distribution (Valtueña et al. 2008). In Spain it is pollinated by three species of Passeriformes (Ortega et al. 2005;Valtueña et al. 2007;Valtueña et al. 2008). It is not uncommon to find ornithophily in Fabaceae, although the family is mainly bee-pollinated (Stone et al 2003;Tandon et al. 2003;Ragusa-Netto 2002;Arroyo 1981). ...
The ornithophilous Anagyris foetida is almost the only shrub in the eastern Mediterranean region that flowers in winter. We assessed the pollination potential of flower visitors by observing pollen loads on trapped animals and contacts with floral parts. Resident and wintering omnivorous bird species, together with honey bees (Apis mellifera), were found to carry the shrub’s pollen throughout its long flowering period in Israel and thus might be legitimate pollinators. Spring-migrating omnivorous birds passing through Israel carried pollen. The Palestine Sunbird (Cinnyris osea) is the only specialist nectarivorous bird in the Middle East. It carries little pollen and is therefore primarily a nectar thief of A. foetida.
... Sugar concentration would be higher if the standing crop experiences a major loss of volume due to evaporation (cf. Valtueña et al., 2007 ). Accordingly, we noted that standing crops in open fl owers that we measured in the aft ernoon tended to have a higher sugar concentration than those measured in the morning. ...
Premise of the study:
Properties of floral nectar have been used to predict if a plant species is pollinated by birds. To see whether winter-flowering plants evolve nectar properties corresponding to bird pollinators, nectar properties of several Camellia species (including the golden-flowered tea), as well as the role of floral visitors as effective pollinators, were examined.
Methods:
Potential pollinators of Camellia petelotii were identified at different times of day and under various weather conditions. A bird exclusion experiment was used to compare the pollination effectiveness of birds and insects. Nectar sugar components (fructose, glucose, and sucrose) from C. petelotii growing wild and another seven Camellia species and 22 additional cultivars (all in cultivation) were examined by high-performance liquid chromatography (HPLC).
Key results:
The sunbird Aethopyga siparaja and honeybees were the most frequent floral visitors to C. petelotii. Honeybee visits were significantly reduced in cloudy/rainy weather. The fruit and seed set of flowers with birds excluded were reduced by 64%, indicating that bird pollination is significant. For the wild populations of C. petelotii, a bagged flower could secrete 157 μL nectar; this nectar has a low sugar concentration (19%) and is sucrose-dominant (87%). The eight Camellia species and 22 cultivars had an average sugar concentration of around 30% and a sucrose concentration of 80%, demonstrating sucrose-dominant nectar in Camellia species.
Conclusions:
The nectar sugar composition of Camellia species was characterized by sucrose dominance. In addition, the large reduction in seed set when birds are excluded in the golden-flowered tea also supports the suggestion that these winter-flowering plants may have evolved with birds as significant pollinators.
... This type of stamens can also attract bees collecting pollen (Miyake et al. 1998). While morphological floral traits tend to be rather constant within a species (Worley and Barrett 2000;Herrera 2001), nectar production and composition and floral scents are physiological traits which depend on the physical conditions of the plant and environmental conditions, which may change over short spatial and temporal scales (Rathcke 1992;Torres and Galetto 1998;Lake and Hughes 1999;Biernaskie and Cartar 2004;Longo and Fischer 2006;Macukanovic-Jocic et al. 2004;Valtuena et al. 2007;Ono et al. 2008). In this way, flowers of E. chiloensis ssp. ...
Flowers that are pollinated both during the day and at night could exhibit two different groups of pollinators and produce two different sets of attractants and rewards. We explored the patterns of emission of flower scents and production of nectar in the cactus Echinopsis chiloensis ssp. chiloensis, in relation to the patterns of activity of its diurnal and nocturnal pollinators. We measured frequency of flower visitors, analyzed floral scents, measured nectar production and sugar concentration, and performed pollination exclusion experiments. Bees were the main visitors at daytime and hawkmoths at nighttime. Diurnal scents were dominated by several compounds that can attract a wide range of pollinators, whereas nocturnal scents were less diverse and were dominated by (E)-nerolidol, a compound eliciting antennal responses in hawkmoths. Nectar volume and sugar concentration at night were similar to those recorded in hawkmoth-pollinated flowers. Daytime nectar volume was higher than those commonly found in bee-pollinated flowers, but similar to those found in flowers pollinated by several pollinators. Daytime sugar concentration was similar to those recorded in bee-pollinated flowers. Flowers of E. chiloensis ssp. chiloensis seem morphologically adapted to hawkmoth pollination, but diurnal and nocturnal pollinators contribute to similar extents to reproductive success. Additionally, diurnal and nocturnal pollinators showed a synergic effect on the product of fruit set and seed set. The results are discussed in terms of the linkage between floral traits and perception abilities and requirements of pollinators.
... The flowers are papilionaceous, pendulous, odorless, yellowish, and borne directly on the trunks (cauliflory) (Ortega-Olivencia et al., 2005). They produce large amounts of very dilute, hexose-rich nectar (Valtueña et al., 2007), and are mainly pollinated by 3 species of passeriforms (Ortega-Olivencia et al., 2005). The fruit is a large pod (45-190 × 12.5-24 mm) with 1-4(8) seeds of large size (9-16 × 6.5-9.5 mm) (Valtueña et al., 2010). ...
... The sites of both populations and the Garden have a Mediterranean climate characterized by rainy, cool winters and dry, hot summers. For more details about the populations, see Ortega-Olivencia et al. (2005) and Valtueña et al. (2007Valtueña et al. ( , 2008a. ...
Anagyris foetida L. has a Mediterranean distribution, but its populations are highly fragmented. This, together with its mixed mating system, puts it at clear risk of being strongly affected by inbreeding depression. To determine the effect of inbreeding depression in this species, hand pollinations were performed during the 2003/04 season in 2 populations from SW Spain to generate selfed and outcrossed seeds. These seeds were divided in 2 sample lots and seedlings were obtained in different years. During the first year of life the first lot showed no differences in vegetative parameters between the 2 populations, so that no differentiation was made between populations in analyzing the results. Reproductive parameters were determined during the first 2 flowering periods in the second sample lot. The results showed inbreeding depression to be manifest in the populations studied throughout the life cycle, adversely affecting the reproductive parameters more than the vegetative parameters, with the male capacity being strongly affected. This shows the need to study the entire life cycle to understand how it is affected by inbreeding depression, and to analyze more than one flowering period in long-lived species. The findings may have important implications for the long-term conservation management of these populations.
... Besides the above mentioned authoritative literature, experts have also relied upon the biology of the reproduction of diff erent taxa of the family Fabaceae , 2000, 2005, Rodríguez-Riaño et al. 1999a, 1999c, 2006, Valtueña et al. 2007, 2008a, 2008b, 2010a, 2010b, 2011 or the genera Drosophyllum (Ortega-Olivencia et al. 1995 or Scrophularia (Ortega-Olivencia and Devesa 1993a, 1993b, Valtueña et al. 2013. Use of these literature resources and long standing experience of the researchers high degree of confidence to the taxonomic identifi cation of the specimens. ...
The herbarium of University of Extremadura (UNEX Herbarium) is formed by 36451 specimens of vascular plants whose main origin is the autonomous region of Extremadura (Spain) and Portugal, although it also contains a smaller number of specimens from different places, including the rest of peninsular Spain, the Baleares Islands, the Macaronesian region (Canary Islands, Madeira and Azores), northwest of Africa (Morocco) and Brazil. 98% of the total records are georeferenced.
It is an active collection in continuous growth. Its data can be accessed through the GBIF data portal at http://data.gbif.org/datasets/resource/255 and http://www.eweb.unex.es/eweb/botanica/herbario/. This paper describes the specimen associated data set of the UNEX Herbarium, with an objective to disseminate the data contained in a data set with potential users, and promote the multiple uses of the data.
... In both cases, the axis of the raceme is triquetrous, with flowers arranged in whorls of three. The flowers produce large amounts of a very dilute and hexose-dominant nectar (Valtueñ a et al. 2007 ). They are mainly pollinated by three passeriforms (two warblers, Sylvia atricapilla L. and Sylvia melanocephala Gmelin, and a chiffchaff, Phylloscopus collybita Vieillot; Ortega-Olivencia et al. 2005). ...
We investigated the role of resource limitation, ovary reserve, and selective abortion in controlling flower and fruit set in the Mediterranean leguminous shrub Anagyris foetida. The removal of bracts, but not that of leaves, reduced the initiation of fruit with respect to controls. The removal of a large proportion (2/3 of the total) of preanthesic inflorescences increased the reproductive capacity of individuals via two different strategies: (1) maintaining the number of fruit-bearing inflorescences, but increasing the number of fruits per inflorescence and seeds per fruit; and (2) increasing the number of fruit-bearing inflorescences, while maintaining fruits/inflorescence and seeds/fruit ratios unchanged. At the level of the inflorescence, most of the ripe fruits were located on the basalmost whorls of the raceme. Within the legume, fertilization was independent of the position of ovules, although the more basal presented lower rates of fertilization than the more central-stylar. The same was observed for the probability of a fertilized ovule to develop into a mature seed.
... In any case, it is possible that the findings of Morrant et al. (2009) for Eucalyptus spathulata are also due to uneven sugar concentration in the nectar of this species. Valtueña et al. (2007) also found two nectar sugar concentrations in flowers of Anagyris fetida; sugar was more concentrated in hanging droplets of this pendulous flower than inside the calyx. Although the concentration trend is opposite to the one we found, possibly because of gravity since flowers of A. patula are not pendulous (A. ...
... The intricacies of nectar production (e.g. Nepi and Stpiczy nska 2008), including nectar heterogeneity within a flower at one point in time (this study; Valtueña et al. 2007), combined with the spatiotemporal variation in visitor assemblages and their effectiveness (e.g. Gómez and Zamora 1999), make ambrosiology a very complex science. ...
Previous research indicated that microcapillary tubes greatly underestimated sugar present in flowers with low nectar volumes, but it was unclear whether tubes missed liquid nectar or whether sugar concentration in nectar they collected did not represent total sugar concentration in a flower. We determined the suitability of microcapillary tubes to estimate the energetic value of Acrotriche patula R.Br. (Ericaceae) nectar from total sugar mass. We collected a standing crop of nectar from individual flowers with microcapillary tubes and subsequently washed the flowers to recover putatively any residual sucrose, glucose, and fructose. We assessed microcapillary nectar volume as a predictor for total sugar mass in a flower by regression analysis, identified the percentage of sugar missed by microcapillary tubes, and compared sugar ratios between microcapillary samples and total nectar. Nectar volume collected with microcapillary tubes cannot be used to predict total nectar sugar contents in a flower. Microcapillary tubes missed 71% of the floral sugar on average, but not a large volume, indicating that sugar is not evenly distributed in a flower’s nectar. Proportions of different sugars did not differ significantly between microcapillary samples and total samples. Animals with different tongue morphologies and feeding behaviours may obtain different energetic rewards from the same flower with low nectar volume. Variation in a flower’s nectar at one point in time is likely to favour the generalisation of pollination systems.
