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This study characterizes the osmophores and corolla traits in 18 species of Bignonieae Dumort., a Bignoniaceae tribe occurring in the Cerrado, a neotropical savanna in Brazil. To detect osmophore distribution, whole, newly opened flowers were immersed in Neutral Red Solution. Samples from the corolla tube and lobes were also fixed and analyzed micromorphologically, anatomically, and histochemically. The osmophores showed six markedly different distribution patterns that were not clearly associated with histological features. In most species, osmophores comprised papillose secretory epidermises and a few layers of subepidermal parenchyma. Starch grains, lipid droplets, and terpenes were detected in osmophores. An ornamented cuticle, cuticular folds, glandular and non-glandular trichomes, raised stomata and epicuticular wax granules are common traits in the species studied and may be useful in determining the taxonomy of the group. We found that 94% of the species visited by bees had papillose epidermises while the single hummingbird-pollinated species presented a flattened epidermis. Variations in osmophore pattern among species visited by bees, including variations within the same plant genus, are novel finding. Additionally, the Bignonieae species visited by bees presented a textured corolla surface, which has been reported as facilitating bee attachment and movement towards the floral resource. Future studies with a greater number of Bignonieae species and more detailed pollinator behavioral assays may help in the interpretation of the variations in corolla traits and functional relationships between flowers and pollinators.

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Lophopterys floribunda is a Neotropical species of Malpighiaceae endemic to Brazil, occurring in both the Amazon and Atlantic Forest. Instead of the typical bi-glandular sepals reported for Neotropical Malpighiaceae, this species presents a single, large gland on the lateral sepals. In addition, ant patrolling was observed at the apex of bracts and bracteoles during fieldwork. Thus, this work aimed to describe the sepalar gland of L. floribunda and other secretory structures in its flowers and inflorescence. Samples of bracts, bracteoles, sepals, petals, and anther were collected and submitted to usual anatomical techniques. Unexpected nectaries at the apex of bracts and bracteoles, not visible to the naked eye, were described and represent a new type of structure for the family due to both their position and size. Mutualistic ants consume the exudate produced by these tiny nectaries, and such structures enable a specific visitation pattern for Lophopterys. Typical epithelial elaiophores occur on the lateral sepals, formed by an invaginated epidermis, which predominantly produce lipid secretion. The petal marginal glands are anatomically similar to the standard type of colleter, which exude mucilaginous substances. The exudate produced by the petal marginal glands was considered to have an additional role of contributing to the maintenance of the closed bud during the beginning of development. The globose epidermal cells containing lipids, proteins, and polysaccharides observed in the connective may be responsible for the typical aroma emitted by these flowers. The diversity of secretory structures reported here has application in both systematic and ecological studies of Malpighiaceae.

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Floral fragrances play an important role in pollinator attraction; they serve as signals for primary rewards such as nectar. The presence of osmophores (a specialized glandular tissue that produces fragrance) in the terrestrial orchid Cyclopogon has only been described in a single species. There are fragrant and odorless species within this genus, but no data on the similarities or differences between them has been published. Here, we present new data on the lip morphology and ultrastructure from eight Cyclopogon species to better understand the mechanisms of pollinator attraction of this group. To achieve this goal, we used light, scanning, and transmission electron microscopy. Five out of eight species presented globular trichomes associated with fragrance production on the abaxial side of the labellum. Together with subepidermal cells, they compose the secretory tissue. Despite the absence of globular trichomes, C. apricus also presented cells associated with fragrance production. Lipid droplets, plastids containing plastoglobuli, and starch were indicators of secretory activity present in these species. This same pattern of fragrance production and release, associated with flower morphology, may result in the restriction of pollination by halictid bees.

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The tribe Schwenckieae (Solanaceae) is characterised by the presence of appendages on the corolla, a diagnostic trait for the group. These appendages constitute a median distal projection of the three-lobed petal and occur in the genera Melananthus and Schwenckia but are absent in Heteranthia. We investigated the micromorphology and anatomical structure of the appendages and lateral petal lobes of Schwenckia americana (two varieties), S. angustifolia, S. curviflora and S. novaveneciana, and Melananthus fasciculatus. We also performed histochemical tests to determine if the appendages are involved in the production of volatiles, acting as a fragrance secretory structure (osmophore). The appendages have a uniseriate epidermis, whose cells store phenolics and lipids. The parenchyma is starch-rich just prior to anthesis in all species studied. The sensory test and anatomical analyses identified scent-secreting tissues, not only in the appendages, but also in the lateral petal lobes, whose cells are papillose with a sculptured surface. The α-naphthol p-phenylenediamine (NADI) reaction detected volatile (essential oils) compounds in S. americana var. americana and S. americana var. angustifolia. We demonstrated the secretory tissues and the production of lipids in the corolla appendages of Schwenckia and Melananthus, which indicate their osmogenic function and probable scent emission to attract pollinators.

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The histochemical studies that search for osmophores differ in the protocol they follow for fixation and discoloration of the samples, and also in the type and number of stains utilized. Despite these differences, all the studies have one point in common: the use of fresh material either collected directly in the field or cultivated in botanical gardens and greenhouses. This is an obvious limitation for the studies of osmophores. Flower parts of herbarium specimens of different dates of collection and different plant families were exposed to variable times and percentages of discoloration agents and under different stains for finding if it is possible to positively test osmophores in this type of material. We obtained positive results discoloring the samples with ethyl alcohol 96° and sodium hypochlorite, and staining with Lugol, Oil Red O, and Neutral Red (TIOFH). A protocol (TIOFH3) for osmophore testing that combines these three stains into a single procedure is proposed.

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Flowers of Anacardiaceae and other Sapindales typically produce nectar, but scent, often associated with a reward for pollinators, has surprisingly been mentioned only rarely for members of the family and order. However, flowers of Anacardium humile and Mangifera indica produce a strong sweet scent. The origin and composition of these floral scents is the subject of this study. Screening of potential osmophores on the petals and investigations of their anatomy were carried out by light, scanning and transmission electron microscopy. The composition of the floral fragrance was characterized by gas chromatography-mass spectrometry. In both species, the base of the adaxial side of each petal revealed specialized secretory epidermal cells which are essentially similar in structure and distinct from all other neighbouring cells. These cells also showed evidence of granulocrine secretory mechanisms and slight specific variations in their subcellular apparatus coinciding with the respective composition of the floral fragrance, predominantly composed of sesquiterpenes in A. humile and monoterpenes in M. indica. This study reports the presence of osmophores for the first time in flowers of Anacardiaceae and confirms the link between the ultrastructural features of their secretory cells and the volatiles produced by the flowers. The flowers of most Sapindales, including Anacardiaceae, are nectariferous. However, the presence of osmophores has only been described for very few genera of Rutaceae and Sapindaceae. Both the occurrence of osmophores and fragrance may have largely been overlooked in Anacardiaceae and Sapindales until now. Further studies are needed to better understand the nature and diversity of the interactions of their nectariferous flowers with their pollinators.

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BACKGROUND: Nectar gain and loss are important flower transitions observed in angiosperms, and are particularly common in orchids. To understand such transitions, the availability of detailed anatomical data and species-level phylogenies are crucial. We investigated the evolution of food deception in Epidendrum, one of the largest orchid genera, using genus phylogeny to map transitions between nectar gain and loss among different clades. Associations between anatomical and histochemical changes and nectar gain and loss were examined using fresh material available from 27 species. The evolution of nectar presence/absence in Epidendrum species was investigated in a phylogenetic framework of 47 species, using one nuclear and five plastid DNA regions available from GenBank and sequenced in this study. RESULTS: The presence or absence of nectar was strongly associated with changes in the inner epidermal tissues of nectaries. Nectar-secreting species have unornamented epidermal tissue, in contrast to the unicellular trichomes found on the epidermis of food deceptive species. Bayesian tests confirmed that transitions occurred preferentially from nectar presence to nectar absence across the Epidendrum phylogeny. In addition, independent nectar loss events were found across the phylogeny, suggesting a lack of constraint for these transitions. CONCLUSIONS: Ornamented nectaries may play an important role in the deceptive pollination strategy by secreting volatile organic compounds and providing tactile stimuli to pollinators. The recurrent and apparently irreversible pattern of nectar loss in Epidendrum suggests that food deception may constitute an alternative evolutionarily stable strategy, as observed in other orchid groups.

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Pollination in Solanum (Solanaceae) species is commonly performed by female bees, which vibrate anthers to extract pollen. Another pollen removal type is by male euglossine bees, milking the anthers when searching for floral scents produced by secretory tissues (osmophorous) at the swollen connective of the anthers of species in the Cyphomandra clade. Some species of this clade, however, are buzz-pollinated and present papillate anthers that should also have secretory activity, a hypothesis here tested. The anthers of Solanum luridifuscescens were fixed at different stages of development and analysed under light microscopy, SEM and TEM. Histochemical tests for the detection of starch and lipids were done. Epidermal cells of the abaxial surface of the anthers were visibly papillose, had large nuclei and dense cytoplasm rich in organelles such as mitochondria and plastids, typical features of secretory tissues. In this site, lipid droplets were detected, concomitantly with starch consumption, compatible with the secretory process in osmophores. No exudate or accumulation of substances was seen on the surface; in agreement with a previous pollination study performed in field conditions, where no pollinators were observed collecting floral scents, only pollen. The histochemical and structural analyses have evidenced the lipidic composition of the secretion, strongly pointing to terpenes as the secreted compounds. Ours findings show that papillae of the anthers have secretory activities that produce lipophilic compounds. This does not result in resources for bees, but could be an evolutionary step to the development of more specialised anthers in the Cyphomandra clade.

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The genus Solanum (family Solanaceae) includes more than 1400 species and has buzz-pollinated flowers with poricidal anthers. The present study aimed to describe the distribution, breeding system and pollination mechanism of Solanum melissarum, a species endemic to Brazil. The study of breeding system was conducted in an urban forest fragment in Jataí, GO. Distribution data were gathered from floristic surveys and digital plant databases. The floral morphology and the pollination mechanism were studied on through field observations and preserved flowers. The breeding system was determined through hand pollination treatments. The species has a distribution only in the Brazilian Atlantic forest coastal, and this study provides the first records of S. melissarum for the state of Goiás. The pendulous flowers have poricidal anthers close to the stigma, with membranous thecae joined by a connective bearing osmophores that attract males of Euglossa cordata bees. As they collect fragrances, the bees press the thecae and pollen is released through a bellows mechanism. Based on the hand-pollination treatments, this species is self-incompatible. Isolated forest fragments may not include enough pollinators to ensure the pollination of plants with specialized systems. However, they are essential for the conservation of species with interesting phytogeographic patterns, such as the vicariance observed in S. melissarum, and for the conservation of regional diversity.(AU)

O gênero Solanum (família Solanaceae) inclui mais de 1400 espécies e apresenta polinização vibrátil e flores com anteras poricidas. O estudo tem o objetivo de descrever a distribuição, sistema reprodutivo e o mecanismo de polinização de Solanum melissarum, uma espécie endêmica do Brasil. O estudo da biologia reprodutiva foi conduzido em um fragmento florestal urbano em Jataí, GO. As informações de distribuição foram obtidas através de trabalhos florísticos e de banco de dados digitais de plantas. A morfologia floral e o mecanismo de polinização foram estudados através de observações diretas em campo e em flores preservadas. O sistema reprodutivo foi determinado a partir de tratamentos de polinização manual. A espécie apresentava distribuição somente na Mata Atlântica litorânea brasileira, e este estudo demonstrou o primeiro registro de S. melissarum para o estado de Goiás. As flores são pendentes e com anteras poricidas próximas ao estigma, com tecas membranosas envoltas por um conectivo giboso com osmóforos que atraem machos da abelha Euglossa cordata. Na coleta de fragrâncias, as abelhas pressionam as tecas e o polén é liberado por um mecanismo de fole. Baseado nos tratamentos de polinização manual, a espécie é autoincompatível. Fragmentos florestais isolados podem não incluir polinizadores suficientes para garantir a polinização das plantas com sistemas especializados, no entanto, eles são essenciais para a conservação de espécies com padrões fitogeográficos interessantes, como o vicariância observada em S. melissarum, e também para a conservação da diversidade regional.(AU)

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Abstract The genus Solanum (family Solanaceae) includes more than 1400 species and has buzz-pollinated flowers with poricidal anthers. The present study aimed to describe the distribution, breeding system and pollination mechanism of Solanum melissarum, a species endemic to Brazil. The study of breeding system was conducted in an urban forest fragment in Jataí, GO. Distribution data were gathered from floristic surveys and digital plant databases. The floral morphology and the pollination mechanism were studied on through field observations and preserved flowers. The breeding system was determined through hand pollination treatments. The species has a distribution only in the Brazilian Atlantic forest coastal, and this study provides the first records of S. melissarum for the state of Goiás. The pendulous flowers have poricidal anthers close to the stigma, with membranous thecae joined by a connective bearing osmophores that attract males of Euglossa cordata bees. As they collect fragrances, the bees press the thecae and pollen is released through a bellows mechanism. Based on the hand-pollination treatments, this species is self-incompatible. Isolated forest fragments may not include enough pollinators to ensure the pollination of plants with specialized systems. However, they are essential for the conservation of species with interesting phytogeographic patterns, such as the vicariance observed in S. melissarum, and for the conservation of regional diversity.

Resumo O gênero Solanum (família Solanaceae) inclui mais de 1400 espécies e apresenta polinização vibrátil e flores com anteras poricidas. O estudo tem o objetivo de descrever a distribuição, sistema reprodutivo e o mecanismo de polinização de Solanum melissarum, uma espécie endêmica do Brasil. O estudo da biologia reprodutiva foi conduzido em um fragmento florestal urbano em Jataí, GO. As informações de distribuição foram obtidas através de trabalhos florísticos e de banco de dados digitais de plantas. A morfologia floral e o mecanismo de polinização foram estudados através de observações diretas em campo e em flores preservadas. O sistema reprodutivo foi determinado a partir de tratamentos de polinização manual. A espécie apresentava distribuição somente na Mata Atlântica litorânea brasileira, e este estudo demonstrou o primeiro registro de S. melissarum para o estado de Goiás. As flores são pendentes e com anteras poricidas próximas ao estigma, com tecas membranosas envoltas por um conectivo giboso com osmóforos que atraem machos da abelha Euglossa cordata. Na coleta de fragrâncias, as abelhas pressionam as tecas e o polén é liberado por um mecanismo de fole. Baseado nos tratamentos de polinização manual, a espécie é autoincompatível. Fragmentos florestais isolados podem não incluir polinizadores suficientes para garantir a polinização das plantas com sistemas especializados, no entanto, eles são essenciais para a conservação de espécies com padrões fitogeográficos interessantes, como o vicariância observada em S. melissarum, e também para a conservação da diversidade regional.

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The constitutive characters of plants can be structural or biochemical and play an important role in their defence against pathogens. Citrus postbloom fruit drop (PFD) caused by Colletotrichum spp. is one of the most important fungal diseases of citrus. The pathogen infects the flowers, leading to premature fruit drop and reducing citrus production. However, flower buds smaller than 8 mm long are usually not infected by Colletotrichum spp. Thus, this study investigated whether there are constitutive mechanisms in flower buds related to Colletotrichum spp. infection. We studied flower buds that were 2, 3, 4, 8, 12 and 15 mm long and petals, after anthesis, of sweet orange 'Valência' using light and scanning electron microscopy and histochemistry. We evaluated the effect of volatile organic compounds (VOCs) in flowers (R-limonene and linalool) on the in vitro growth of Colletotrichum acutatum. We found that the arrangement of the epidermal papillae in the petal primordia, the occurrence of prismatic crystals and the distribution of oil glands are the main differences between buds smaller than 8 mm and buds 8-15 mm long. Osmophores at the tips of petals produced and accumulated phenols, terpenes and lipophilic compounds. Flower buds smaller than 8 mm long have constitutive structural and biochemical barriers to Colletotrichum spp. infection. In addition, this is the first time that osmophores have been reported in citrus. Our study shows that natural terpenes of Citrus flowers inhibit the fungal growth in vitro, highlighting the potential use of terpenes for the chemical control of PFD in citrus.

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Scent glands, or osmophores, are predominantly floral secretory structures that secrete volatile substances during anthesis, and therefore act in interactions with pollinators. The Leguminosae family, despite being the third largest angiosperm family, with a wide geographical distribution and diversity of habits, morphology and pollinators, has been ignored with respect to these glands. Thus, we localised and characterised the sites of fragrance production and release in flowers of legumes, in which scent plays an important role in pollination, and also tested whether there are relationships between the structure of the scent gland and the pollinator habit: diurnal or nocturnal. Flowers in pre-anthesis and anthesis of 12 legume species were collected and analysed using immersion in neutral red, olfactory tests and anatomical studies (light and scanning electron microscopy). The main production site of floral scent is the perianth, especially the petals. The scent glands are distributed in a restricted way in Caesalpinia pulcherrima, Anadenanthera peregrina, Inga edulis and Parkia pendula, constituting mesophilic osmophores, and in a diffuse way in Bauhinia rufa, Hymenaea courbaril, Erythrostemon gilliesii, Poincianella pluviosa, Pterodon pubescens, Platycyamus regnellii, Mucuna urens and Tipuana tipu. The glands are comprised of cells of the epidermis and mesophyll that secrete mainly terpenes, nitrogen compounds and phenols. Relationships between the presence of osmophores and type of anthesis (diurnal and nocturnal) and the pollinator were not found. Our data on scent glands in Leguminosae are original and detail the type of diffuse release, which has been very poorly studied.

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