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Nuptial and extranuptial nectaries are involved in interactions with different animal functional groups. Nectar traits involved in pollination mutualisms are well known. However, we know little about those traits involved in other mutualisms, such as ant-plant interactions, especially when both types of nectaries are in the same plant organ, the flower. Here we investigated if when two types of nectaries are exploited by distinct functional groups of floral visitors, even being within the same plant organ, the nectar secreted presents distinct features that fit animal requirements. We compared nectar secretion dynamics, floral visitors and nectar chemical composition of both nuptial and extranuptial nectaries in natural populations of the liana Amphilophium mansoanum (Bignoniaceae). For that we characterized nectar sugar, amino acid and specialized metabolite composition by high-performance liquid chromatography. Nuptial nectaries were visited by three medium- and large-sized bee species and extranuptial nectaries were visited mainly by ants, but also by cockroaches, wasps and flies. Nuptial and extranuptial nectar differed regarding volume, concentration, milligrams of sugars per flower and secretion dynamics. Nuptial nectar was sucrose-dominated, with high amounts of γ-aminobutyric acid and ß-aminobutyric acid and with theophylline-like alkaloid, which were all exclusive of nuptial nectar. Whereas extranuptial nectar was hexose-rich, had a richer and less variable amino acid chemical profile, with high amounts of serine and alanine amino acids and with higher amounts of the specialized metabolite tyramine. The nectar traits from nuptial and extranuptial nectaries differ in energy amount and nutritional value, as well as in neuroactive specialized metabolites. These differences seem to match floral visitors' requirements, since they exclusively consume one of the two nectar types and may be exerting selective pressures on the composition of the respective resources of interest.

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Mabea fistulifera, a species pollinated mainly by diurnal and nocturnal vertebrates, presents pendulous inflorescences with approximately 70 pairs of nuptial nectaries (NNs). These NNs exude voluminous nectar drops that defy gravity, remaining exposed at the inflorescence for more than a day. We aimed to investigate the NN secretory process and the unique nectar presentation of M. fistulifera. NNs and their exudate were collected at different secretory stages and submitted to structural studies and chemical analysis. The epidermis is devoid of stomata and constitutes the main site of synthesis for non-sugar metabolites found on nectar and nectar-coating film. Nectary parenchyma presents few small starch grains, and vascular strands are distributed until the nectary parenchyma cells close to the epidermis. Vascular tissues at the nectary parenchyma seem to provide sugar and water for the nectar. A film composed of lipids, alkaloids, and proteins covers the nectar drops. The film guarantees the nectar offering for several hours, as it minimizes water loss and prevents falls by gravitational action. The release of large nectar drops is intrinsically linked to the NN anatomical traits and the exudate composition. Low sugar concentration and predominance of hexoses in M. fistulifera nectar are essential for maintaining nectar exudation for many hours, which results in the visitation of a broad spectrum of pollinators.

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Barkeria scandens and B. whartoniana are endangered, endemic taxa from Mexico. They are epiphytes adapted to dry habitats. Since these plants are xerophytic, their flowers were investigated for structural adaptations to nectar secretion. The flowers of both species are structurally similar, and contrary to most claims for the genus, have functional floral nectaries comprising a nectary chamber and a narrow tubular cuniculus. Nectar is present in both these structures, and contains sugars and lipid-like compounds. The nectary tissue is composed of a single-layered epidermis overlying 1-2 layers of subepidermal secretory parenchyma. The outer tangential wall of the epidermal cells is thick and multi-layered, whereas the cuticle, which often shows blistering, is lamellate and possesses micro-channels. Lipid-like material occurs both between the microfibrils of the cell wall and in the micro-channels. Robust secretory tissue, thick cell walls, and lipid-like nectar components limit nectar evaporation. Moreover, the rigidity of the nectary potentially makes it possible for red-flowered B. scandens to switch from entomophily to ornithophily.

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MAIN CONCLUSION: The cytological changes underlying the formation of an apoplasmic barrier in the multi-layered extrafloral nectaries of Citharexylum myrianthum are compatible with the synthesis, transport and deposition of suberin. In terms of ontogenesis and function, the intermediate layers of these nectaries are homologous with the stalks of nectar-secreting trichomes. Anticlinal cell wall impregnations are common in trichomatic nectaries and their functions as endodermis-like barriers have been discussed because of possible direct effects on the nectary physiology, mainly in the nectar secretion and resorption. However, the cytological events linked to nectary wall impregnations remain little explored. This study documents the ontogenesis and the fine structure of the EFN cells, and cytological events linked to the wall impregnations of multi-layered extrafloral nectaries (EFNs) in Citharexylum myrianthum Cham. (Verbenaceae). EFNs are patelliform, and differentiated into (a) a multicellular foot, which is compound in structure and vascularised with phloem strands, (b) a bi-layered intermediate region with thickened cell walls and (c) a single-layered secretory region with palisade-like cells. EFNs are protodermal in origin, starting with a single protodermal cell and ending with the complex, multi-layered structure. The cell wall impregnations first appear in the very young EFN and increase towards maturity. Lipid patches (assumed to be suberin) are deposited on the inner faces of the primary walls, first along the anticlinal walls and then extend to the periclinal walls. On both walls, plasmodesmata remain apparently intact during the maturation of the EFNs. In the peripheral cytoplasm there are abundant polymorphic plastids, well-developed Golgi bodies often close to rough endoplasmic reticulum profiles, mitochondria and polyribosomes. Cytological events linked to the wall impregnations are consistent with suberin synthesis, transport and deposition. Our findings offer new insights into the structure-properties of specialised nectary cell walls and so should contribute to our knowledge of the physiological and protective roles of this structure in nectar glands.

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PREMISE: Floral rewards are essential in understanding floral function and evolution of the relationships between flowers and pollinators. Whether sugars are present in stigmatic exudates in Anthurium and whether it has floral nectaries have remained controversial because of the scarcity of structural studies. To solve these questions, we investigated the floral anatomy of A. andraeanum to elucidate whether (1) tepals are secretory organs, (2) tepals possess a structurally recognizable nectary, and (3) tepalar secretion differs from stigmatic secretion. METHODS: Floral structure was assessed through light and electron microscopy of samples of immature, pistillate, and staminate flowers. The dynamics of the starch reserve was investigated using histochemical tests, and the sugar content in the floral exudates was assessed using thin-layer chromatography. RESULTS: Sugar analysis did not detect sucrose, glucose, or fructose in stigmatic secretions, but confirmed their presence in tepalar secretions. Stigmatic secretion was produced by secretory stigmatic papillae; tepalar exudates were produced by nonvascularized nectaries in the apex of tepals. These nectaries were characterized by modified stomata and cells with cytoplasm rich in organelles, and a high content of calcium oxalate crystals. CONCLUSIONS: Our results showed for the first time nectaries on tepals and true nectar secretion for A. andraeanum. Stigmatic secretion appears to be a distinct substance, and its often-reported sugar content seems to be a result of sample contamination. Nectar and stigmatic secretions have been often mistaken in other Anthurium species and deserve a revision for this genus.

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The presence of nectarless flowers in nectariferous plants is a widespread phenomenon in angiosperms. However, the frequency and distribution of nectarless flowers in natural populations, and the transition from nectariferous to nectarless flowers are poorly known. Variation in nectar production may affect mutualism stability, since energetic resource availability influences pollinators' foraging behavior. Here, we described the spatial and temporal nectar production patterns of Jacaranda oxyphylla, a bee-pollinated species that naturally presents nectarless flowers. Additionally, we compared nectariferous and nectarless floral disks in order to identify histological, subcellular and chemical changes that accompanied the loss of nectar production ability. For that we used standard methods for light and transmission electron microscopy, and gas chromatography coupled to mass spectrometry for chemical analyses. We verified that 47% of flowers did not produce nectar during the whole flower lifespan (nectarless flowers). We also observed remarkable inter-plant variation, with individuals having only nectarless flowers, others only nectariferous ones and most of them showing different proportions of both flower types, with variable nectar volumes (3-21 µl). Additionally, among nectariferous flowers, we registered two distinct rhythms of nectar production. 'Early' flowers produced nectar from 0 to 24 h, and 'late' flowers produced nectar from 24 to 48 h of anthesis. Although disks from nectariferous and nectarless flowers displayed similar histological organization, they differed strongly at subcellular level. Nectariferous ('early' and 'late') flowers exhibited a cellular apparatus typical of nectar secretion, while nectarless flowers exhibited osmophoric features. We found three aliphatic and one aromatic compound(s) that were detected in both the headspace of flowers and the disks of nectarless flowers, but not the disks of nectariferous flowers Although the remarkable variation in nectar availability may discourage pollinator visits, nectarless flowers might compensate it by producing volatile compounds that can be part of floral scent, acting as chemical attractants. Thus, nectarless flowers may be helping to maintain pollination in this scenario of trophic resource supply scarcity. We suggest that J. oxyphylla can be transitioning from a nectar-based pollination system to another resource-based or even to a deceit mechanism of pollination.

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Relationships among floral biology, floral micromorphology and pollinator behaviour in bird-pollinated orchids are important issues to understand the evolution of the huge flower diversity within Orchidaceae. We aimed to investigate floral mechanisms underlying the interaction with pollinators in two hummingbird-pollinated orchids occurring in the Atlantic forest. We assessed floral biology, nectar traits, nectary and column micromorphologies, breeding systems and pollinators. In both species, nectar is secreted by lip calli through spaces between the medial lamellar surfaces of epidermal cells. Such a form of floral nectar secretion has not been previously described. Both species present functional protandry and are self-compatible yet pollinator-dependent. Fruit set in hand-pollination experiments was more than twice that under natural conditions, evidencing pollen limitation. The absence of fruit set in interspecific crosses suggests the existence of post-pollination barriers between these sympatric co-flowering species. In Elleanthus brasiliensis, fruits resulting from cross-pollination and natural conditions were heavier than those resulting from self-pollination, suggesting advantages to cross-pollination. Hummingbirds pollinated both species, which share at least one pollinator species. Species differences in floral morphologies led to distinct pollination mechanisms. In E. brasiliensis, attachment of pollinarium to the hummingbird bill occurs through a lever apparatus formed by an appendage in the column, another novelty to our knowledge of orchid pollination. In E. crinipes, pollinarium attachment occurs by simple contact with the bill during insertion into the flower tube, which fits tightly around it. The novelties described here illustrate the overlooked richness in ecology and morphophysiology in Orchidaceae.

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Nectaries occur widely in Convolvulaceae. These structures remain little studied despite their possible importance in plant-animal interactions. In this paper, we sought to describe the structure and ultrastructure of the receptacular nectaries (RNs) of Ipomoea cairica, together with the dynamics of nectar secretion. Samples of floral buds, flowers at anthesis and immature fruits were collected, fixed and processed using routine methods for light, scanning and transmission electron microscopy. Circadian starch dynamics were determined through starch measurements on nectary sections. The secretion samples were subjected to thin layer chromatography. RNs of I. cairica were cryptic, having patches of nectar-secreting trichomes, subglandular parenchyma cells and thick-walled cells delimiting the nectary aperture. The glandular trichomes were peltate type and had typical ultrastructural features related to nectar secretion. The nectar is composed of sucrose, fructose and glucose. Nectar secretion was observed in young floral buds and continued as the flower developed, lasting until the fruit matured. The starch content of the subglandular tissue showed circadian variation, increasing during the day and decreasing at night. The plastids were distinct in different portions of the nectary. The continuous day-night secretory pattern of the RNs of I. cairica is associated with pre-nectar source circadian changes in which the starch acts as a buffer, ensuring uninterrupted nectar secretion. This circadian variation may be present in other extrafloral nectaries and be responsible for full daytime secretion. We conclude that sampling time is relevant in ultrastructural studies of dynamic extranuptial nectaries that undergo various changes throughout the day.

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The pollination of Passiflora coccinea by the hummingbird Phaethornis superciliosus was studied in Central Amazon, Brazil. We hypothesized that a greater nectar secretion rate (NSR) increases the pollination success of single flowers through Ph. superciliosus visiting behavior. For control flowers, NSR was an increasing function of flower base diameter (FBD). The total number of Ph. superciliosus probes per flower was an increasing function of FBD. Additionally, deposition of pollen on stigmas increased with the cumulative number of Ph. superciliosus probes. Our results show that larger P. coccinea flowers secrete nectar at higher rates, are probed more times during each hummingbird visit and are more successful at pollination. This seems to be the first non-manipulative study describing such an effect of NSR on the pollination of single flowers in nature.

Estudamos a polinização de Passiflora coccinea por beija-flores Phaethornis superciliosus na Amazônia Central, Brasil. Nossa hipótese é que maiores taxas de secreção de néctar (TSN) aumentam o sucesso da polinização de flores individuais através do comportamento de visitas de Ph. superciliosus. Para flores controladas, a TSN foi uma função positiva do diâmetro da base da flor (DBF). O número total de visitas de Ph. superciliosus por flor foi uma função positiva do DBF. Adicionalmente, a deposição de pólen sobre os estigmas aumentou com o aumento do número acumulado de visitas de Ph. superciliosus. Nossos resultados indicam que flores maiores de P. coccinea secretam néctar em taxas mais altas, são visitadas mais vezes pelos beija-flores, e apresentam maior sucesso de polinização. Este parece ser o primeiro estudo não-manipulativo que descreve este efeito da TSN sobre o sucesso de polinização de flores individuais na natureza.

RESUMO

The pollination of Passiflora coccinea by the hummingbird Phaethornis superciliosus was studied in Central Amazon, Brazil. We hypothesized that a greater nectar secretion rate (NSR) increases the pollination success of single flowers through Ph. superciliosus visiting behavior. For control flowers, NSR was an increasing function of flower base diameter (FBD). The total number of Ph. superciliosus probes per flower was an increasing function of FBD. Additionally, deposition of pollen on stigmas increased with the cumulative number of Ph. superciliosus probes. Our results show that larger P. coccinea flowers secrete nectar at higher rates, are probed more times during each hummingbird visit and are more successful at pollination. This seems to be the first non-manipulative study describing such an effect of NSR on the pollination of single flowers in nature.

Estudamos a polinização de Passiflora coccinea por beija-flores Phaethornis superciliosus na Amazônia Central, Brasil. Nossa hipótese é que maiores taxas de secreção de néctar (TSN) aumentam o sucesso da polinização de flores individuais através do comportamento de visitas de Ph. superciliosus. Para flores controladas, a TSN foi uma função positiva do diâmetro da base da flor (DBF). O número total de visitas de Ph. superciliosus por flor foi uma função positiva do DBF. Adicionalmente, a deposição de pólen sobre os estigmas aumentou com o aumento do número acumulado de visitas de Ph. superciliosus. Nossos resultados indicam que flores maiores de P. coccinea secretam néctar em taxas mais altas, são visitadas mais vezes pelos beija-flores, e apresentam maior sucesso de polinização. Este parece ser o primeiro estudo não-manipulativo que descreve este efeito da TSN sobre o sucesso de polinização de flores individuais na natureza.

RESUMO

Fertile ramets of bumblebee-pollinated Alstroemeria aurea, a clonal perennial native to the temperate forests of the southern Andes, produce single terminal inflorescences that may bear two or more temporally non-overlapping whorls of flowers. While fruit set is commonly high (>80%) among early-opening flowers, it is usually low (<20%) among late-opening flowers within ramets. Using flowering ramets with two whorls of flowers, we examined experimentally the following related hypotheses. First, late flowers act as a reserve of ovaries, increasing their likelihood of setting seed when early fruits abort due to either pollen or resource limitation. Second, where early fruit abortion has occurred, plants may actively ensure pollination of late flowers by increasing their attractants and rewards. In a natural population, we simulated (1) lack of pollen deposition in early flowers, by excising their stigmas just before receptivity, and (2) resource limitation, by removing all the leaves from an experimental flowering ramet. Treatments were applied to individual ramets according to a 2 × 2 factorial design. We found that when early flowers failed to set fruit due to stigma excision, nectar secretion and particularly pollen receipt strongly increased in late flowers. Higher pollen deposition contributed significantly to the observed five-fold increase in seed output of late flowers. Fruit and seed set from early flowers were more negatively affected by defoliation than that from late flowers. Defoliation did not interfere with a ramet's capacity to increase late reproductive output when early reproduction failed. These results support the assertion that late flowers act as a reserve of ovaries helping a plant to cope with an unpredictable environment. These results also suggest that plants may actively increase pollinator visitation by opportunistically increasing flower rewards.