Nectar and oleiferous trichomes as floral attractants in Bulbophyllum saltatorium Lindl. (Orchidaceae)

Although many Orchidaceae have deceit flowers that produce no reward, the most common reward, when present, is nectar. Bulbophyllum, however, is unusual in that the labellar secretions of most species investigated to date lack sugars, and, therefore, cannot be considered true nectar. The African species Bulbophyllum saltatorium is an exception in that it produces not only nectar but also possesses specialized, capitate oleiferous trichomes. The nectary of B. saltatorium is borne on the labellum and is represented by a deep, narrow, median longitudinal groove, having a small aperture, and flanked by trichomes. Isodiametric epidermal cells lining this groove secrete nectar which collects both in the groove and on the surface of the labellum. As well as a nectary, the labellum of B. saltatorium also bears three types of unicellular trichomes: the longest trichomes are borne distally and abaxially; the marginal ones form a rim around the entire labellum, and finally, massive, capitate trichomes occur proximally and adaxially. These are oleiferous, containing large quantities of oil which might function as precursors of volatile components of fragrance or provide a food-reward. To the best of our knowledge, this is the first time for such oleiferous trichomes to be described for Bulbophyllum. Therefore, apart from their color and markings, flowers of this species are able to attract pollinators in at least two, possibly three ways: food-reward in the form of nectar; fragrance; and possibly food-rewards in the form of food-hairs

The structure of the spur nectary in Dendrobium finisterrae Schltr. (Dendrobiinae, Orchidaceae)

To date, the structure of the nectary spur of Dendrobium finisterrae has not been studied in detail, and the present paper compares the structural organization of the floral nectary in this species with the spurs of other taxa. The nectary spur of D. finisterrae was examined by means of light microscopy (LM), scanning electron microscopy (SEM), and transmission electron microscopy (TEM). It is composed of a single layer of secretory epidermis and several layers of small and compactly arranged subepidermal secretory cells. The secretory cells have thick cellulosic cell walls with primary pits. The secretory tissue is supplied by vascular bundles that run beneath in ground parenchyma and are additionally surrounded by strands of sclerenchymatous fibers. The flowers of the investigated species displayed morphological features characteristic of bee-pollinated taxa, as they are zygomorphic, creamy-green coloured with evident nectar guides. They also emit a weak but nice scent. However, they possess some characters attributed to bird-pollinated flowers such as a short, massive nectary spur and collenchymatous secretory tissue that closely resembles the one found in the nectaries of certain species that are thought to be bird-pollinated. This similarity in anatomical organization of the nectary, regardless of geographical distribution and phylogeny, strongly indicates convergence and appears to be related to pollinator-driven selection

Floral micromorphology of the Australian carnivorous bladderwort Utricularia dunlopii, a putative pseudocopulatory species

Flowers of sexually deceptive taxa generally possess a set of morphological and physiological characters that mimic their insect pollinators. These characters often include a specific insect-like floral configuration, together with scent glands osmophores) that produce fragrances which chemically resemble insect sex pheromones. Furthermore, these flowers tend not to produce pollinator food rewards. According to some authors, flowers of the Australian bladderwort Utricularia dunlopii (and species of the Utricularia capilliflora complex) resemble insects, and pollination perhaps occurs by pseudocopulation. The aims of this paper are to compare the structure and distribution of floral glandular trichomes in the Australian carnivorous plant U. dunlopii with those of closely related species assigned to the same section and to discuss their putative function. Floral tissues of U. dunlopii P. Taylor, Utricularia paulinae Lowrie, Utricularia dichotoma Labill. and Utricularia uniflora R.Br. (section Pleiochasia) were investigated using light microscopy, scanning electron microscopy, transmission electron microscopy and histochemistry. In U. dunlopii, two long, erect, filiform appendages arising from the upper lip of the corolla, together with three arising from the lower lip, bear numerous glandular trichomes that may function as osmophores. In other species, such as U. uniflora and U. paulinae, glandular papillae on the corolla palate may also function as osmophores. The floral anatomical and morphological organisation of U. dunlopii differs from that of the other investigated species, indicating that its insect pollinators are also likely to differ. Morphological and ultrastructural observations, while generally contributing to our understanding of the flower of U. dunlopii, do not refute the possibility that pollination here may occur by pseudocopulation. Further field-based investigations, however, are now necessary to test this hypothesis

Structure of the cuniculus nectary in Brassavola flagellaris Barb. Rodr. (Laeliinae Benth., Orchidaceae)

To date, the structure of the cuniculus nectary has not been studied in detail. Furthermore, the secretory mechanism of such nectaries has not been investigated. The present paper describes, for the first time, the structural organization and ultrastructure of the cuniculus nectary in the moth-pollinated orchid Brassavola flagellaris Barb. Rodr. This tubular structure is situated between the perianth tube and ovary and, in its possession of thick, cellulose cell walls, resembles the nectary of ornithophilous taxa. The presence of large secretory vesicles that fuse with the plasmalemma indicate that granulocrine nectar secretion occurs in this species. The lumen of the cuniculus is lined with unicellular hairs. However, the cuticle overlying the whole epidermal surface lining the lumen (both glabrous and pubescent regions) was coated with nectar residues and became distended and cracked, indicating that this entire tissue is probably involved in nectar secretion

Spatiotemporal variation in the pollination systems of a supergeneralist plant: is Angelica sylvestris (Apiaceae) locally adapted to its most effective pollinators?

Background and Aims: In terms of pollination systems, umbellifers (plants of the carrot family, Apiaceae) are regarded as generalists, since their (usually dichogamous) flowers are visited by a wide range of insects representing several taxonomic orders. However, recent analyses of insect effectiveness revealed that these plants may be pollinated effectively by a narrow assemblage of insect visitors. Of particular interest was whether populations of an umbellifer species varied in pollinator assemblages and whether this could lead to local specialization of the pollination system. We also explored whether variation in pollinator assemblages was associated with variation in floral traits, and whether this variation influences reproductive output. Methods: The focus was on Angelica sylvestris, a common European species visited by a taxonomically diverse insect assemblage. In three populations, located along an ~700-km transect, over three growth seasons insect visitors were identified, their effectiveness was assessed by surveying pollen loads present on the insect body, insect activity on umbels, nectar and scent composition was studied, and transplantation experiments were performed. Key Results: The populations investigated in this study differed in their nectar and scent profiles and, despite the similar taxonomic composition of insect visitor assemblages, were effectively pollinated by disparate pollinator morphogroups, i.e. flies and beetles. Although this suggested local adaptations to the most effective pollinators, analyses of body pollen loads and behaviour on umbels demonstrated functional equivalency of the visitor morphogroups, which is probably related to the fact that A. sylvestris bears few ovules per flower. The transplantation experiments confirmed that reproductive success was not related to the source of experimental plants and that the insects do not exhibit preferences towards local genotypes. Conclusions: Angelica sylvestris is morphologically well adapted to ecological generalization, and there is little evidence that the surveyed populations represent distinct pollination ecotypes. Most likely, the observed variation in floral characters can be interpreted as 'adaptive wandering'. Specialization in this family seems possible only under very special circumstances, for example when the pollinator community comprises insect visitor groups that clearly differ in their pollination capacity (e.g. due to differences in their functional morphology) and/or have different perceptional biases (e.g. for colour or scent). However, the barrier to the evolution of morphological adaptations resulting in the fine-tuning of the flower towards particular pollinator types may arise from the architectural constraints on the floral bauplan that make umbellifers so uniform in their floral displays and so successful in attracting large numbers of pollinators

Floral ultrastructure of two Brazilian aquatic-epiphytic bladderworts : Utricularia cornigera Studnička and U. nelumbifolia Gardner (Lentibulariaceae)

Utricularia cornigera and Utricularia nelumbifolia are giant, aquatic-epiphytic species of carnivorous bladderwort from southeastern Brazil that grow in the central ‘urns’ of bromeliads. Both species have large, colourful flowers. The main aim of our study is to ascertain whether the prominent floral palate of U. cornigera and U. nelumbifolia functions as an unguentarius-i.e. an organ that bears osmophores. Floral tissues of both species were investigated using light microscopy, scanning electron microscopy, transmission electron microscopy and histochemistry. Floral palates of U. cornigera and U. nelumbifolia provide clear visual signals for pollinating insects. In both species, the palate possesses diverse micro-morphology, comprising unicellular, conical to villiform papillae and multicellular, uniseriate, glandular trichomes that frequently display terminal branching. The most characteristic ultrastructural feature of these papillae was the presence of relatively large, polymorphic plastids (chromoplasts) containing many plastoglobuli. Similar plastids are known to occur in the fragrance-producing (osmophores) and oil-producing (elaiophores) tissues of several orchid species. Thus, these palate papillae may play a key role in providing the olfactory stimulus for the attraction of insect pollinators. Nectariferous trichomes were observed in the floral spurs of both species, and in U. nelumbifolia, free nectar was also recorded. The location, micro-morphology, anatomy and ultrastructure of the floral palate of the two species investigated may thus indicate that the palate functions as an unguentarius. Furthermore, the flowers of these taxa, like those of U. reniformis, have features consistent with bee pollination

Flower palate structure of the aquatic bladderworts Utricularia bremii Heer and U. minor L. from section Utricularia (Lentibulariaceae)

There is an enormous diversity in the structure of the flower palate of the carnivorous rootless genus Utricularia. This study aims to examine the structure of the palates in Utricularia bremii Heer and U. minor L of the Utricularia sect. Utricularia, which have a glandular palate type. In both species, the palate has only one type of glandular trichomes. Because of the occurrence of cell wall ingrowths in its glandular cells, any exudation may be transported via eccrinous secretion. It was proposed that the palate trichomes of the examined species act as scent glands and that the palate may play a role as an unguentarium. Both U. bremii and U. minor are of an open flower type. Thus, U. bremii and U. minor flowers can be penetrated by small, weak insects, which then easily have access to their generative structure. Small Hymenoptera (member of families Mymaridae and Braconidae) were observed as flower visitors of the male-sterile species Utricularia bremii

Nectar-Secreting and Nectarless Epidendrum: Structure of the Inner Floral Spur

Epidendrum, the largest genus of Neotropical orchids, contains both nectar-secreting and nectarless species. Here, we compare the fine structure of the inner floral spur, termed the cuniculus, in nectariferous (E. difforme, E. nocturnum,E. porpax, E. rigidum, E. vesicatum) and seemingly nectarless (E. capricornu, E. ciliare, E. criniferum, E. pseudepidendrum, E. radicans, E. xanthoianthinum) species. This is the first time for such a detailed investigation of cuniculus structure to be undertaken for Epidendrum. Our aim was to characterize features indicative of secretory activity and to ascertain whether flowers presumed to be nectarless produce alternative pollinator food-rewards. The cuniculus is formed by fusion of the basal part of the labellum and column and extends alongside the ovary and transmitting tract. Our study indicates that all investigated species produce nectar or nectar-like secretion to varying degrees, and no alternative pollinator food-rewards were observed. Even though macroscopic investigation of presumed rewardless species failed to reveal the presence of secretion within the cuniculus, close observations of the cells lining the cuniculus by LM, SEM, and TEM revealed the presence of cuticular blisters and surface material. Moreover, the similarity of both the thick tangential cell walls (with the exception of E. vesicatum) and organelle complement of cuniculus epidermal cells in both copiously nectariferous species and those producing only small quantities of surface secretion confirmed the presence of secretory activity in species generally regarded to be rewardless. The secretory character was particularly obvious in the cells of the cuniculus of E. nocturnum, but also in E. ciliare, E. radicans and E. xanthoianthinum, since electron-dense cytoplasm and mitochondria, ER and secretory vesicles were abundant. Furthermore, cell wall protuberances occurred in E. nocturnum, which was indicative of intense transmembrane transport. This investigation highlights the need to examine more closely whether Epidendrum spp. considered to lack food-rewards based solely on macroscopic examination really are rewardless and deceptive

Nectar trichome structure of aquatic bladderworts from the section Utricularia (Lentibulariaceae) with observation of flower visitors and pollinators

In Utricularia, the flower spur is a nectary and in this organ, nectar is produced and stored. This study aimed to examine the structure of the nectary trichomes in four Utricularia species (Utricularia vulgaris L., U. australis R.Br., U. bremii Heer and U. foliosa L.) from the generic section Utricularia. We have investigated whether species with different spur morphology had similar spur anatomy and nectary trichome structure. In Utricularia flowers, nectar is produced by spur capitate trichomes (sessile or stalked). Our results showed that regardless of the various spur morphology, trichomes have similar architecture and ultrastructure. Head cells of these trichomes are transfer cells with an eccrine nectar secretion. Examined species differed in the micromorphology of papillae in spurs. The fly Eristalis tenax was found to be a pollinator of U. vulgaris. Small Halictidae bees seem to be pollinators of U. foliosa

Floral micromorphology of the bird-pollinated carnivorous plant species Utricularia menziesii R.Br. (Lentibulariaceae)

Made available in DSpace on 2019-10-06T16:15:22Z (GMT). No. of bitstreams: 0 Previous issue date: 2019-01-01• Background and Aims Bird pollination is rare among species in the genus Utricularia, and has evolved independently in two lineages of this genus. In Western Australia, the Western Spinebill, Acanthorhynchus superciliosus, visits flowers of Utricularia menziesii (section Pleiochasia: subgenus Polypompholyx). This study aimed to examine the micromorphology of U. menziesii flowers to assess traits that might be linked to its pollination strategy. • Methods Light microscopy, histochemistry and scanning electron microscopy were used. Nectar sugar composition was analysed using high-performance liquid chromatography. • Key Results The flowers of U. menziesii fulfil many criteria that characterize bird-pollinated flowers: red colour, a large, tough nectary spur that can withstand contact with a hard beak, lack of visual nectar guides and fragrance. Trichomes at the palate and throat may act as tactile signals. Spur nectary trichomes did not form clearly visible patches, but were more frequently distributed along vascular bundles, and were small and sessile. Each trichome comprised a single basal cell, a unicellular short pedestal cell (barrier cell) and a multicelled head. These trichomes were much smaller than those of the U. vulgaris allies. Hexose-dominated nectar was detected in flower spurs. Fructose and glucose were present in equal quantities (43 ± 3.6 and 42 ± 3.6 g L-1). Sucrose was only detected in one sample, essentially at the limit of detection for the method used. This type of nectar is common in flowers pollinated by passerine perching birds. • Conclusions The architecture of nectary trichomes in U. menziesii was similar to that of capitate trichomes of insect-pollinated species in this genus; thus, the most important specializations to bird pollination were flower colour (red), and both spur shape and size modification. Bird pollination is probably a recent innovation in the genus Utricularia, subgenus Polypompholyx, and is likely to have evolved from bee-pollinated ancestors.Department of Plant Cytology and Embryology Jagiellonian University in Kraków, 9 Gronostajowa St.Faculty of Biology University of Warsaw Botanic Garden, Al. Ujazdowskie 4Department of Animal Histology and Embryology University of Silesia in Katowice, 9 Bankowa St.School of Biological Sciences University of Western Australia (M084), 35 Stirling HighwayDepartamento de Biologia Aplicada à Agropecuária Faculdade de Ciências Agrárias E Veterinárias Universidade Estadual Paulista (Unesp)Faculty of Science School of Biological Sciences University of AdelaideUnit of Botany and Plant Physiology Institute of Plant Biology and Biotechnology University of Agriculture in Kraków, 29 Listopada 54 StreetDepartamento de Biologia Aplicada à Agropecuária Faculdade de Ciências Agrárias E Veterinárias Universidade Estadual Paulista (Unesp