The need for weeds: Man-made, non-cropped habitats complement crops and natural habitats in providing honey bees and bumble bees with pollen resources.

In Europe, honey bees and bumble bees are among the most important pollinators, and there is a growing interest in understanding the effects of floral resource availability on their survival. Yet, to date, data on nectar and pollen supplies available to bees in agricultural landscapes are still scarce. In this paper, we quantify species-, habitat- and landscape-scale pollen production in the Lublin Upland, SE Poland. The production per unit area was highest (mean = 2.2-2.6 g/m2) in non-forest woody vegetation, field margins and fallows, whilst significantly lower pollen amounts were shown to be available in road verges and railway embankments (mean = 1.3-1.6 g/m2). At landscape scale, natural and semi-natural areas (forests and meadows/pastures) offered ca. 44% of the total pollen resources during the year. Relatively high amounts of pollen (ca. 35% of the year-round total pollen resources) were from winter rape, but this resource was short-term. Man-made, non-cropped habitats added only ca. 18% of the total pollen mass offered for pollinators during flowering season. However, they provided 66-99% of pollen resources available from July to October. There exists an imbalance in the availability of pollen resources throughout the year. Hence, a diversity of natural, semi-natural and man-made, non-cropped areas is required to support the seasonal continuity of pollen resources for pollinators in an agricultural landscape. Efforts should be made to secure habitat heterogeneity to enhance the flower diversity and continual pollen availability for pollinators.

Phenology and production of pollen, nectar, and sugar in 1612 plant species from various environments.

To predict the quantity and quality of food available to pollinators in various landscapes over time, it is necessary to collect detailed data on the pollen, nectar, and sugar production per unit area and the flowering phenology of plants. Similar data are needed to estimate the contribution of plants to the functioning of food webs via the flow of energy and nutrients through the soil-plant-nectar/pollen-consumer pathway. Current knowledge on this topic is fragmented. This database represents the first compilation of data on the various food resources produced by 1612 plant species belonging to 755 genera and 133 families, including crop plants and wild plants, annuals and perennials, animal- and wind-pollinated plants, and weeds and trees growing in different ecosystems under various environmental conditions. The data set consists of 103 parameters related to the traits of plant species and geographical and environmental factors, allowing for precise calculations of the amounts of nectar, pollen, and energy provided by plants and available to consumers in the considered flora or ecosystem on a daily basis throughout the year. These parameters, gathered by us and extracted from the available literature, describe pollen, nectar, and sugar production (where applicable, in mass, volume, and concentration units), honey yield, the timing and duration of flowering, flower longevity, number of plants and flowers per unit area, weather conditions (temperature and precipitation), geographical location, landscape, and syntaxonomy. The data were obtained from various, mostly European, pedoclimatic zones, and the majority of the data were available for plant species and communities present in Central Europe, especially in Poland, where research on floral resources has a long tradition. These data are representative of the whole continent and may be used as a reference for plant communities occurring on continents other than Europe since the database allows for the consideration of differences in the production of resources by a single plant species growing in different communities. This data set provides a unique opportunity to test hypotheses related to the functioning of food webs, nutrient cycling, plant ecology, and pollinator ecology and conservation. The data are released under a CC-BY-NC-SA license, and this paper must be properly cited when using the database.

Unravelling the dependence of a wild bee on floral diversity and composition using a feeding experiment.

We investigated nutrition as a potential mechanism underlying the link between floral diversity/composition and wild bee performance. The health, resilience, and fitness of bees may be limited by a lack of nutritionally balanced larval food (pollen), influencing the entire population, even if adults are not limited nutritionally by the availability and quality of their food (mainly nectar). We hypothesized that the nutritional quality of bee larval food is indirectly connected to the species diversity of pollen provisions and is directly driven by the pollen species composition. Therefore, the accessibility of specific, nutritionally desirable key plant species for larvae might promote bee populations. Using a fully controlled feeding experiment, we simulated different pollen resources that could be available to bees in various environments, reflecting potential changes in floral species diversity and composition that could be caused by landscape changes. Suboptimal concentrations of certain nutrients in pollen produced by specific plant species resulted in reduced bee fitness. The negative effects were alleviated when scarce nutrients were added to these pollen diets. The scarcity of specific nutrients was associated with certain plant species but not with plant diversity. Thus, one of the mechanisms underlying the decreased fitness of wild bees in homogenous landscapes may be nutritional imbalance, i.e., the scarcity of specific nutrients associated with the presence of certain plant species and not with species diversity in pollen provisions eaten by larvae. Accordingly, we provide a conceptual representation of how the floral species composition and diversity can impact bee populations by affecting fitness-related life history traits. Additionally, we suggest that mixes of 'bee-friendly' plants used to improve the nutritional base for wild bees should be composed considering the local flora to supplement bees with vital nutrients that are scarce in the considered environment.

Habitat heterogeneity helps to mitigate pollinator nectar sugar deficit and discontinuity in an agricultural landscape.

The scarcity of floral resources and their seasonal discontinuity are considered as major factors for pollinator decline in intensified agricultural landscapes worldwide. The consequences are detrimental for the stability of the environment and ecosystems. Here, we quantified the production of nectar sugars in plant species occurring in man-made, non-cropped areas (non-forest woody vegetation, road verges, railway embankments, field margins, fallow areas) of an agricultural landscape in SE Poland. We also assessed changes in the availability of sugar resources both in space (habitat and landscape scales) and in time (throughout the flowering season), and checked to what extent the sugar demands of honeybees and bumblebees are met at the landscape scale. At landscape-level, 37.6% of the available sugar resources are produced in man-made, non-cropped habitats, while 32.6% and 15.0% of sugars derive from winter rape crops and forest vegetation, respectively. Nectar sugar supplies vary greatly between man-made, non-cropped habitat types/sub-types. These areas are characterized by a high richness of nectar-producing species. However, a predominant role in total sugar resources is ascribable to a few species. Strong fluctuations in nectar resources are recorded throughout the flowering season. March and June are periods with food shortages. Abundant nectar sugars are generally found in April-May, mainly due to the mass flowering of nectar-yielding species in the forests, meadows/pastures and orchards/rapeseed crops. Heterogeneity of man-made, non-cropped habitats is essential to support the supply of July-October nectar sugars for honeybees and bumblebees. Reduced flowering in man-made non-cropped habitats can generate serious food deficiencies, as from summer towards the end of the flowering season >90% of sugars are provided by the flora of these areas. Therefore, highly nectar-yielding plant species that flower during periods of expected food shortages should be a priority for conservation and restoration programs.

Spikelet movements, anther extrusion and pollen production in wheat cultivars with contrasting tendencies to cleistogamy.

BACKGROUND: Cleistogamic flowers are a main barrier in pollen dispersal for cross-pollination necessary in wheat hybrid breeding. The aim of our study was to gain new knowledge on the biology of wheat flowering, in particular on the differences between the cleisto- and chasmogamic forms which has certainly cognitive significance, but it can also be used in practice when seeking a female and male ideotypes for cross hybridization. RESULTS: We characterized the most significant features defining the flowering specificity in two wheat cultivars with contrasting tendency to cleistogamy: Piko (chasmogamous) and Dacanto (cleistogamous). In the field observations we assessed diurnal pattern of anther extrusion and anther extrusion capacity. For the first time we adapted the time lapse method for measuring kinetics of the spikelet movement and 3-D image correlation technique for the non-invasive measurements of potential deformations of the spikelet lemmas. We found that the two cultivars differ in the potential of pollen dispersion for-cross-pollination and in the spikelet kinetics. We also described some anatomical traits that can have potential functional role in floret opening. None of the cultivars showed any symptoms of lemma surface deformation. CONCLUSIONS: The cleistogamic and chasmogamic wheat cultivars differ significantly in the potential for pollen dispersion for cross-pollination, which is mainly related to anther extrusion capacity. Although none of these features differentiated the cultivars clearly, we assume, based on spikelet kinetics and the lack of lemmas surface deformation, that the water transport and turgor of cells is essential for the floret opening and anther extrusion in wheat. The search for parental ideotype should be supported by marker assisted selection, e.g. based of polymorphisms in genes related to aquaporin biosynthesis.

Asteraceae species as potential environmental factors of allergy.

The statistics from Europe and the USA have proven a high risk for skin diseases associated with plant contact. Therefore, plant-induced dermatitis is of increasing attention in dermatology. The focus of this paper was to present the current knowledge on aspects of contact allergy related to Asteraceae (Compositae) species. The Asteraceae family is one of the largest in the world with members across all continents. The PubMed/Medline databases have been searched. The Asteraceae representatives consist of diverse secondary metabolites, which exhibit various advantageous effects in humans. In particular, sesquiterpene lactones (SLs) may cause sensitization resulting in skin irritation and inflammation. In this study, we tried to reveal the allergenic potential of several Asteraceae species. The Asteraceae-related allergy symptoms involve eczema, hay fever, asthma, or even anaphylaxis. Furthermore, the evidence of severe cross-reactivity with food and pollen allergens (PFS) in patients sensitive to Asteraceae allergens have been announced. Further identification and characterization of secondary metabolites and possible allergens in Asteraceae are necessary for the better understanding of Asteraceae-related immune response. The Asteraceae allergy screening panel (the SL mix and the Compositae mix of five plant species) is a promising tool to improve allergy diagnostics and therapy.

Micromorphological and histochemical attributes of flowers and floral reward in Linaria vulgaris (Plantaginaceae).

The self-incompatible flowers of Linaria vulgaris have developed a range of mechanisms for attraction of insect visitors/pollinators and deterrence of ineffective pollinators and herbivores. These adaptive traits include the flower size and symmetry, the presence of a spur as a "secondary nectar presenter," olfactory (secondary metabolites) and sensual (scent, flower color, nectar guide-contrasting palate) signals, and floral rewards, i.e. pollen and nectar. Histochemical tests revealed that the floral glandular trichomes produced essential oils and flavonoids, and pollen grains contained flavonoids, terpenoids, and steroids, which play a role of olfactory attractants/repellents. The nectary gland is disc-shaped and located at the base of the ovary. Nectar is secreted through numerous modified stomata. Nectar secretion began in the bud stage and lasted to the end of anthesis. The amount of produced nectar depended on the flower age and ranged from 0.21 to 3.95 mg/flower (mean = 1.51 mg). The concentration of sugars in the nectar reached up to 57.0%. Both the nectar amount and sugar concentration demonstrated a significant year and population effect. Pollen production was variable between the years of the study. On average, a single flower of L. vulgaris produced 0.31 mg of pollen. The spectrum of insect visitors in the flowers of L. vulgaris differed significantly between populations. In the urban site, Bombus terrestris and Apis mellifera were the most common visitors, while a considerable number of visits of wasps and syrphid flies were noted in the rural site.

Validation of floral food resources for pollinators in agricultural landscape in SE Poland.

BACKGROUND: Proper management of bee pastures is considered an important activity for diversity of pollinators and conservation of the population size. In the present study, the floral composition and diversity, flowering spectrum and availability of food resources in natural and man-made habitats in four rural municipalities in Lublin Upland, SE Poland were evaluated. The connection between pollinator-friendly biotopes was also determined and some suggestions are made on creation/supplementation of bee pastures adapted to the arable land structure in eastern Poland (mean acreage 7.65 ha). RESULTS: Forage species richness (S) differed among vegetation types: the highest S value was found in field margins, while mixed forests represented the lowest species richness (mean ± standard deviation = 37.8 ± 9.5 vs 16.5 ± 2.3). In all habitats, the most abundant flowering was recorded in May. Then the blooming abundance decreased until a complete decline at the end of summer. The mean distance between pollinator-friendly biotopes is 2.4 ± 1.53 km, which is out of the flying range/foraging distance of most pollinators. CONCLUSION: Shortages of nectar and pollen resources in the agricultural landscape of SE Poland are evident. Therefore it is assumed that the landscape structure requires support in terms of food niches for pollinators and creation/supplementation of bee pastures is necessary, e.g. six to nine patches of 0.025-0.3 ha each within an area of 100 ha. © 2017 Society of Chemical Industry.

Floral nectary, nectar production dynamics and chemical composition in five nocturnal Oenothera species (Onagraceae) in relation to floral visitors.

Main conclusion The floral nectars were sucrose-dominant; however, nectar protein and amino acid contents differed, indicating that composition of nitrogenous compounds may vary considerably even between closely related plant species, irrespectively of nectary structure. Numerous zoophilous plants attract their pollinators by offering floral nectar; an aqueous solution produced by specialized secretory tissues, known as floral nectaries. Although many papers on nectaries and nectar already exist, there has been a little research into the structure of nectaries and/or nectar production and composition in species belonging to the same genus. To redress this imbalance, we sought, in the present paper, to describe the floral nectary, nectar production, and nectar composition in five nocturnal Oenothera species with respect to their floral visitors. The structure of nectaries was similar for all the species investigated, and comprised the epidermis (with nectarostomata), numerous layers of nectary parenchyma, and subsecretory parenchyma. Anthesis for a single flower was short (ca. 10-12 h), and flowers lasted only one night. The release of floral nectar commenced at the bud stage (approx. 4 h before anthesis) and nectar was available to pollinators until petal closure. Nectar concentration was relatively low (ca. 27%) and the nectar was sucrose-dominant, and composed mainly of sucrose, glucose and fructose. The protein content of the nectar was also relatively low (on average, 0.31 µg ml-1). Nevertheless, a great variety of amino acids, including both protein and non-protein types, was detected in the nectar profile of the investigated taxa. We noted both diurnal and nocturnal generalist, opportunistic floral insect visitors.

Ecological stoichiometry of the honeybee: Pollen diversity and adequate species composition are needed to mitigate limitations imposed on the growth and development of bees by pollen quality.

The least understood aspects of the nutritional needs of bees are the elemental composition of pollen and the bees' need for a stoichiometrically balanced diet containing the required proportions of nutrients. Reduced plant diversity has been proposed as an indirect factor responsible for the pollinator crisis. We suggest stoichiometric mismatch resulting from a nutritionally unbalanced diet as a potential direct factor. The concentrations and stoichiometric ratios of C, N, S, P, K, Na, Ca, Mg, Fe, Zn, Mn, and Cu were studied in the bodies of honeybees of various castes and sexes and in the nectar and pollen of various plant species. A literature review of the elemental composition of pollen was performed. We identified possible co-limitations of bee growth and development resulting mainly from the scarcity of Na, S, Cu, P and K, and possibly Zn and N, in pollen. Particular castes and sexes face specific limitations. Concentrations of potentially limiting elements in pollen revealed high taxonomic diversity. High floral diversity may be necessary to maintain populations of pollen eaters. Single-species crop plantations, even if these species are rich in nectar and pollen, might limit bee growth and development, not allowing for gathering nutrients in adequate proportions. However, particular plant species may play greater roles than others in balancing honeybee diets. Therefore, we suggest specific plant species that may (1) ensure optimal growth and production of individuals by producing pollen that is exceptionally well balanced stoichiometrically (e.g., clover) or (2) prevent growth and development of honeybees by producing pollen that is extremely unbalanced for bees (e.g., sunflower). Since pollen is generally poor in Na, this element must be supplemented using "dirty water". Nectar cannot supplement the diet with limiting elements. Stoichiometric mismatch should be considered in intervention strategies aimed at improving the nutritional base for bees.

Biological and therapeutic properties of bee pollen: a review.

Natural products, including bee products, are particularly appreciated by consumers and are used for therapeutic purposes as alternative drugs. However, it is not known whether treatments with bee products are safe and how to minimise the health risks of such products. Among others, bee pollen is a natural honeybee product promoted as a valuable source of nourishing substances and energy. The health-enhancing value of bee pollen is expected due to the wide range of secondary plant metabolites (tocopherol, niacin, thiamine, biotin and folic acid, polyphenols, carotenoid pigments, phytosterols), besides enzymes and co-enzymes, contained in bee pollen. The promising reports on the antioxidant, anti-inflammatory, anticariogenic antibacterial, antifungicidal, hepatoprotective, anti-atherosclerotic, immune enhancing potential require long-term and large cohort clinical studies. The main difficulty in the application of bee pollen in modern phytomedicine is related to the wide species-specific variation in its composition. Therefore, the variations may differently contribute to bee-pollen properties and biological activity and thus in therapeutic effects. In principle, we can unequivocally recommend bee pollen as a valuable dietary supplement. Although the bee-pollen components have potential bioactive and therapeutic properties, extensive research is required before bee pollen can be used in therapy. © 2016 Society of Chemical Industry.

Floral nectar production and carbohydrate composition and the structure of receptacular nectaries in the invasive plant Bunias orientalis L. (Brassicaceae).

The data relating to the nectaries and nectar secretion in invasive Brassicacean taxa are scarce. In the present paper, the nectar production and nectar carbohydrate composition as well as the morphology, anatomy and ultrastructure of the floral nectaries in Bunias orientalis were investigated. Nectary glands were examined using light, fluorescence, scanning electron and transmission electron microscopy. The quantities of nectar produced by flowers and total sugar mass in nectar were relatively low. Total nectar carbohydrate production per 10 flowers averaged 0.3 mg. Nectar contained exclusively glucose (G) and fructose (F) with overall G/F ratio greater than 1. The flowers of B. orientalis have four nectaries placed at the base of the ovary. The nectarium is intermediate between two nectary types: the lateral and median nectary type (lateral and median glands stay separated) and the annular nectary type (both nectaries are united into one). Both pairs of glands represent photosynthetic type and consist of epidermis and glandular tissue. However, they differ in their shape, size, secretory activity, dimensions of epidermal and parenchyma cells, thickness of secretory parenchyma, phloem supply, presence of modified stomata and cuticle ornamentation. The cells of nectaries contain dense cytoplasm, plastids with starch grains and numerous mitochondria. Companion cells of phloem lack cell wall ingrowths. The ultrastructure of secretory cells indicates an eccrine mechanism of secretion. Nectar is exuded throughout modified stomata.