Osmophores and petal surface traits in Bignonieae species.

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.

To each their own! Nectar plasticity within a flower mediates distinct ecological interactions.

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.

Phenological data of a Neotropical savanna community.

Savanna plant species commonly have different adaptive mechanisms in response to fire. In this biome, phenology is a functional trait characterizing the responses of plant communities to fire. The database presented here provides phenological data on 95 angiosperm species, in plots with natural vegetation and/or in burned plots. We used 10 plots (5 × 5 m) installed in "campo cerrado" physiognomy, in the Santa Bárbara Ecological Station, located in the municipality of Águas de Santa Bárbara, São Paulo state, southeastern Brazil. Half of these plots was burned in Aug/2013 and the other half was kept intact as a control. For one year (Sep/2013 to Aug/2014), we collected monthly data on the presence of sprouting (new branches or new unexpanded leaves), flowers (flower buds and flowers at anthesis) and fruits (immature and mature) in all angiosperm individuals present in the plots. This phenological data can support other studies on these sampled species, involving different aspects of their ecology, and on the conservation of this type of vegetation and management plans in relation to the prescription of fire.

Florivory and Pollination Intersection: Changes in Floral Trait Expression Do Not Discourage Hummingbird Pollination.

Many flowers are fed on by florivores, but we know little about if and how feeding on flowers affects their visual and chemical advertisement and nectar resource, which could disrupt pollination. Here, we investigated if damages caused by florivores compromise a Neotropical hummingbird pollination system, by modifying the floral advertisements and the nectar resource. We surveyed natural florivory levels and patterns, examined short-term local effects of floral damages caused by the most common florivore, a caterpillar, on floral outline, intra-floral colour pattern and floral scent, as well as on the amount of nectar. Following, we experimentally tested if the most severe florivory pattern affected hummingbird pollination. The feeding activity of the most common florivore did not alter the intra-floral colour pattern, floral scent, and nectar volume, but changed the corolla outline. However, this change did not affect hummingbird pollination. Despite visual floral cues being important for foraging in hummingbirds, our results emphasise that changes in the corolla outline had a neutral effect on pollination, allowing the maintenance of florivore-plant-pollinator systems without detriment to any partner.

Floral UV Features of Plant Species From a Neotropical Savanna.

Despite the wide interest in flower colours, only after the end of the nineteenth-century studies started to comprise floral UV reflection, which is invisible to humans but visible to the major groups of pollinators. Many flowers and inflorescences display colour patterns, an important signal for pollinators, promoted by the presence of at least two different colours within flowers or inflorescences, including colours in the UV waveband. For Neotropical savanna plant species, we characterised floral UV features using UV-photography and reflectance measurements. We tested (i) whether floral UV features were constrained by their shared ancestry, (ii) whether floral UV features were associated with pollinators, and (iii) whether floral UV features were associated with floral traits mediating these interactions, including floral resource, type of attraction unit and presence/absence of non-UV colour patterns. Of 80 plant species, ca. 70% were UV-patternless, most of them UV-absorbing. Approximately 30% presented one of three types of UV-patterns: bullseye, contrasting corolla markings oriented toward floral resources or contrasting reproductive structures, which were all considered as floral guides. Floral UV features were phylogenetically constrained and were associated with pollinators, floral resources and attraction unit, but not with non-UV colour patterns. UV-patternless flowers were associated with most of the pollination systems, while UV-patterned flowers were mainly associated with bee-pollination. UV-absorbing flowers comprised the only category with hawkmoth- and butterfly-pollinated flowers, and a high percentage of hummingbird-pollinated species. Nocturnal pollinated species were also commonly UV-absorbing, except for one UV-reflecting bat-pollinated species and one beetle-pollinated species with UV-reflecting stigmas. All types of floral UV features were associated with nectar; however, flowers with contrasting reproductive structures were mainly associated with pollen. There was an association between UV-absorbing species and the presence of inflorescences and intermediate attraction units. Our results evince that phylogenetic relatedness can constraint floral UV features' diversification, but combinations of evolutionary and ecological processes may be expected in this scenario.

Epicormic bud protection traits vary along a latitudinal gradient in a neotropical savanna.

Regrowth via production of epicormic shoots is an important strategy for many woody plants after environmental disturbances such as fire, drought, and herbivory. Populations spreading across a broad latitudinal gradient offer opportunities to investigate if essential traits vary with heterogenous environmental conditions, such as in savanna ecosystems. This information can help us predict plant responses to climate change. Here, we evaluated if epicormic bud protection traits varied among populations of three focal savanna species (Miconia albicans, Solanum lycocarpum, and Zeyheria montana) that have a wide distribution and grow under variable climatic conditions. We randomly sampled 225 individuals over five spatially independent sites (7°, 10°, 15°, 18°, and 24° S) in Brazil, totaling 15 individuals per species per area. We analyzed anatomical transverse sections of five buds per species per area to assess the relative area occupied by crystal and phenolic idioblasts, the thickness of the trichome boundary layer, and to test if these traits were associated with climatic conditions. The buds were protected by cataphylls and composed of a variable number of undeveloped leaves enveloping the shoot apex. For M. albicans, we found an association between maximum temperature and both phenolic idioblasts and trichome boundary layer, but no association with crystal idioblasts. In S. lycocarpum, only the trichome boundary layer was associated with maximum temperature plus high radiation. Z. montana showed no variation. Combination of two or more traits can lead to the development of adaptative strategies to different climatic conditions. We present for the first time an analysis of epicormic bud traits in plant populations occurring in an extensive latitudinal gradient and shed light on how maximum temperature is associated with these traits, contributing to a better understanding of plant resprouting capabilities in widespread savanna plant species.

I've been robbed! - Can changes in floral traits discourage bee pollination?

Some floral visitors collect nectar by piercing flower external whorls, acting as nectar robbers. They leave robbery vestiges, which can cause changes in floral characteristics, including physical and chemical signals that may influence flower recognition by pollinators. If pollinating bees associate these changes with absence or reduction in nectar volume, they can avoid these flowers, negatively affecting pollination. We aimed to investigate the effect of robbery on primary and secondary attractants. Additionally, we experimentally investigated if the visual signs present in robbed flowers affect the bee pollination of this plant species by discouraging pollinator visits. This study was performed in a very common pollinator-plant-cheaters system comprised by a bee-pollinated Bignoniaceae species and a nectar-robber bee that lands on the corolla tube and makes slits at its base during the nectar robbery. We experimentally isolated the effect of nectar consumption by this nectar-robber and investigated if the slits caused by the nectar-robbers affected the floral scent emission. In addition, we experimentally evaluated the effect of visual signs (slits) associated to the nectar robbery and the effect of nectar depletion on the pollination of Jacaranda caroba (Bignoniaceae). The robbers visited around 75% of the flowers throughout the day and removed significant amounts of nectar from them. However, the damages the robbers cause did not affect floral scent emission and we did not verify significant differences on pollen deposition neither when comparing flowers with slits and control nor when comparing flowers with and without nectar. We showed that even though nectar-robbers visually honestly signal the robbery and deplete high amounts of nectar, they did not affect pollinator visitation. These results showed that presumably antagonistic interactions might in fact not be so.

Nectar Replaced by Volatile Secretion: A Potential New Role for Nectarless Flowers in a Bee-Pollinated Plant Species.

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.

Colleters in Rubiaceae from forest and savanna: the link between secretion and environment.

This study aims to investigate colleters' secretory function, on cellular level, in Rubiaceae species from contrasting environments looking to explore the association between secretion and environment. We collected samples from eight species of Rubiaceae growing in forest and savanna having standard-type colleters with diverse histochemistry (hydrophilic, lipophilic and mixed secretions) and processed for both conventional and cytochemical study under transmission electron microscopy (TEM). The standard colleters, although similar in morphology and anatomy, exhibited marked differences on cellular level, especially in the abundance and topology of Golgi bodies, endoplasmic reticulum and plastids when comparing forest and savanna species. These differences were clearly aligned with the chemical nature of the secretions they produce, with predominance of hydrophilic secretions in forest species and lipophilic or mixed secretions in savanna species. The combination of methods in electron microscopy revealed the sites of synthesis and intracellular compartmentation of substances, the mechanisms of their secretion from the protoplast and confirmed the involvement of the outer walls of the epithelial cells in the elimination of exudates to the gland surface. Our study suggests a potential environment-associated plasticity of the secretory cells of standard-type colleters in modulating their secretory function performance.

Morphology, secretion composition, and ecological aspects of stipular colleters in Rubiaceae species from tropical forest and savanna.

Colleters are secretory structures that produce and release mucilage or a mucilage-resin mixture protecting meristems and young structures against desiccation, herbivores, and pathogens. The secretions may vary in colleters of same or different types, indicating that the functionality of colleters may be more specific than previously thought. In this study, we compared 17 Rubiaceae species from savanna and forest environment focusing on colleter secretions and its ecological role. First, we evaluated the morphology, distribution, and histochemistry of stipular colleters using light and scanning electron microscopy. Additionally, we investigated the phenology, microclimate, and the proportion of damaged apices in the savanna and forest species. We recorded standard-type colleters, variable in distribution and size, in 14 of the 17 studied species. The secretion varied from predominantly hydrophilic, mixed to predominantly lipophilic. During the budding period, secretion covered the vegetative apices. Savanna species had a prevalence of lipid secretion in habitats with higher luminosity, which had a lower proportion of damaged apices. In contrast, forest species occurred in habitats with lower luminosity and had a higher proportion of damaged apices, in general with the absence of lipids in the colleters. These results highlight that colleters with similar morphology clearly differed in secretions among species, especially between species from savanna and forest, in which the colleters appear potentially associated with protection against irradiation in savanna, but not in the forest environment.

Trichome structure and evolution in Neotropical lianas.

BACKGROUND AND AIMS: Trichomes are epidermal outgrowths generally associated with protection against herbivores and/or desiccation that are widely distributed from ferns to angiosperms. Patterns of topological variation and morphological evolution of trichomes are still scarce in the literature, preventing valid comparisons across taxa. This study integrates detailed morphoanatomical data and the evolutionary history of the tribe Bignonieae (Bignoniaceae) in order to gain a better understanding of current diversity and evolution of trichome types. METHODS: Two sampling schemes were used to characterize trichome types: (1) macromorphological characterization of all 105 species currently included in Bignonieae; and (2) micromorphological characterization of 16 selected species. Individual trichome morphotypes were coded as binary in each vegetative plant part, and trichome density and size were coded as multistate. Ancestral character state reconstructions were conducted using maximum likelihood (ML) assumptions. KEY RESULTS: Two main functional trichome categories were found: non-glandular and glandular. In glandular trichomes, three morphotypes were recognized: peltate (Pg), stipitate (Sg) and patelliform/cupular (P/Cg) trichomes. Non-glandular trichomes were uniseriate, uni- or multicellular and simple or branched. Pg and P/Cg trichomes were multicellular and non-vascularized with three clearly distinct cell layers. Sg trichomes were multicellular, uniseriate and long-stalked. ML ancestral character state reconstructions suggested that the most recent common ancestor (MRCA) of Bignonieae probably had non-glandular, Pg and P/Cg trichomes, with each trichome type presenting alternative histories of appearance on the different plant parts. For example, the MRCA of Bignonieae probably had non-glandular trichomes on the stems, prophylls, petiole, petiolule and leaflet veins while P/Cg trichomes were restricted to leaflet blades. Sg trichomes were not present in the MRCA of Bignonieae independently of the position of these trichomes. These trichomes had at least eight independent origins in tribe. CONCLUSIONS: The patterns of trichome evolution indicate that most morphotypes are probably homologous in Bignonieae and could be treated under the same name based on its morphological similarity and common evolutionary history, in spite of the plethora of names that have been previously designated in the literature. The trichome descriptions presented here will facilitate comparisons across taxa, allowing inferences on the relationsthips between trichome variants and future studies about their functional properties.

Functional aspects of floral nectar secretion of Ananas ananassoides, an ornithophilous bromeliad from the Brazilian savanna.

BACKGROUND AND AIMS: Several members of Bromeliaceae show adaptations for hummingbird pollination in the Neotropics; however, the relationships between floral structure, nectar production, pollination and pollinators are poorly understood. The main goal of this study was to analyse the functional aspects of nectar secretion related to interaction with pollinators by evaluating floral biology, cellular and sub-cellular anatomy of the septal nectary and nectar composition of Ananas ananassoides, including an experimental approach to nectar dynamics. METHODS: Observations on floral anthesis and visitors were conducted in a population of A. ananassoides in the Brazilian savanna. Nectary samples were processed using standard methods for light and transmission electron microscopy. The main metabolites in nectary tissue were detected via histochemistry. Sugar composition was analysed by high-performance liquid chromatography (HPLC). The accumulated nectar was determined from bagged flowers ('unvisited'), and floral response to repeated nectar removal was evaluated in an experimental design simulating multiple visits by pollinators to the same flowers ('visited') over the course of anthesis. KEY RESULTS: The hummingbirds Hylocharis chrysura and Thalurania glaucopis were the most frequent pollinators. The interlocular septal nectary, composed of three lenticular canals, extends from the ovary base to the style base. It consists of a secretory epithelium and nectary parenchyma rich in starch grains, which are hydrolysed during nectar secretion. The median volume of nectar in recently opened 'unvisited' flowers was 27·0 µL, with a mean (sucrose-dominated) sugar concentration of 30·5 %. Anthesis lasts approx. 11 h, and nectar secretion begins before sunrise. In 'visited' flowers (experimentally emptied every hour) the nectar total production per flower was significantly higher than in the 'unvisited' flowers (control) in terms of volume (t = 4·94, P = 0·0001) and mass of sugar (t = 2·95, P = 0·007), and the concentration was significantly lower (t = 8·04, P = 0·0001). CONCLUSIONS: The data suggest that the total production of floral nectar in A. ananassoides is linked to the pollinators' activity and that the rapid renewal of nectar is related to the nectary morphological features.

Structuring software anthropometric variables on CD Rom as a facilitator of the process of design of work situations.

The environment where everyday activities are developed, these should be appropriate and the individual who will execute them, for greater efficiency of their work and even if you feel safe, comfortable and satisfied when interacting with that space, expending less energy to this interaction. Given this context, this study aims to structure a software with the results obtained in the field, from anthropometric measure to subsidize the furniture industry in manufacturing of furniture, definition work situations, considering the different anthropometric measurements made between 2001 and 2010, using data from this sample of adults aged 18 to 65 years old and children between 6 and 11 years old. It was convenient to make the software with a site, only instead of staying on the Web has been recorded on CD Rom. Tests with the prototype allow navigation through the structure of the software. The data needed to implement the remaining modules were also raised. Issues related to system layout and usability of the interface also were not considered, because it is an initial prototype.

Determining anthropometric variables as a tool in the preparation of furniture and designs of interior spaces: the case of children 6 to 11 years old of Vicosa, State of Minas Gerais, Brazil.

The adequacy of facilities and the individual securities in their different age groups is importance to ensure greater functionality to them, allowing full development of daily activities. For this to occur more efficiently it is necessary the use of ergonomics which can ensure more comfort and safety for end users of products and spaces. The present study aimed to measure body dimensions of a representative sample of children aged 6 to 11 years old, children of graduate and pos graduate students, faculty and staff of the Federal University of Vicosa and also residents of the city of Vicosa, State of Minas Gerais, Brazil, coming from different municipalities of State of Minas Gerais, to organize a database that will provide the furniture industry, anthropometric variables more appropriate to design products for both the leisure activities, and for the school sector. To realize this research we used the methodology proposed by the authors Panero and Zelnik, based on samples distributed in six age groups, and providing a measurement of 10 variables. By applying the methodology to the field was possible to compare the observed data, with the tables of the aforementioned authors. The main results revealed a significant variation of the 10 variables analyzed, and it is believed that this variation could lead to possible flaws in the designs of products that use the data from these authors. The completion of the study provided data on Vicosa considered more appropriate for the design of products and environments for the population of the study, considering age and region, of Brazil (State of Minas Gerais) and it is believed that the future may expand to the Brazilian population, with the progress of study of this nature.

Pollination biology of Jacaranda oxyphylla with an emphasis on staminode function.

BACKGROUND AND AIMS: Bignoniaceae is a Neotropical family with >100 genera, only two of which, Jacaranda and Digomphia, have a developed staminode. Jacaranda oxyphylla, whose flowers possess a conspicuous glandular staminode, is a zoophilous cerrado species. Here, the composition of the secretion of the glandular trichome and the influence of the staminode on the pollination biology and reproductive success of J. oxyphylla were studied. METHODS: The floral morphology, pollen viability, stigma receptivity, nectar volume and nectar concentration were studied. Compatibility system experiments were performed and floral visitors were observed and identified. Experiments comparing the effect of staminode presence and absence on pollen removal and pollen deposition efficiency were conducted in open-pollinated flowers. Histochemistry, thin-layer chromatography (TLC) and gas chromatography coupled to flame ionization detection (GC-FID) analyses were performed to determine the main chemical components of the staminode's glandular trichome secretion. KEY RESULTS: Flower anthesis lasted 2 d and, despite the low frequency of flower visitation, pollination seemed to be effected mainly by medium-sized Eulaema nigrita and Bombus morio bees, by the small bee Exomalopsis fulvofasciata and occasionally by hummingbirds. Small bees belonging to the genera Ceratina, Augochlora and Trigona were frequent visitors, collecting pollen. Jacaranda oxyphylla is predominantly allogamous. Staminode removal resulted in fewer pollen grains deposited on stigmas but did not affect total pollen removal. The secretion of capitate glandular trichome occurs continually; the main chemical compounds detected histochemically were phenolic and terpenoid (essential oils and resins). Monoterpene cineole, pentacyclic triterpenes and steroids were identified by TLC and GC-FID. CONCLUSIONS: The staminode of J. oxyphyllla is multifunctional and its importance for female reproductive success was attributed mainly to the secretion produced by capitate glandular trichomes. This secretion is involved in complex chemical interactions with pollinating bees, including the solitary bees Euglossini. These bees are common pollinators of various species of Jacaranda.

Ovary peltate trichomes of Zeyheria montana (Bignoniaceae): developmental ultrastructure and secretion in relation to function.

BACKGROUND AND AIMS: Nectar production in the Bignoniaceae species lacking a nectariferous functional disc is ascribed to trichomatic glands around the ovary base and/or on the inner corolla wall. Nevertheless, knowledge about the secretion and function of these glands is very incomplete. The purpose of this paper is to study, from a developmental viewpoint, the ultrastructure, histochemistry and secretory process of the peltate trichomes on the ovary of Zeyheria montana, a species in the Bignoniaceae which has a rudimentary disc. METHODS: Samples of the gynoecium at various developmental stages were fixed and processed for light and electron microscopy. Histochemistry and cytochemistry tests were performed to examine the chemical composition of exudates. Thin layer chromatography was used to determine the presence of alkaloids and terpenes in gynoecium and fruit extracts, and in fresh nectar stored in the nectar chamber. KEY RESULTS: Peltate trichomes at different developmental stages appear side by side from floral budding up to pre-dispersal fruit. Large plastids with an extensive internal membrane system consisting of tubules filled with lipophilic material, abundant smooth endoplasmic reticulum, few Golgi bodies, lipophilic deposits in the smooth endoplasmic reticulum and mitochondria, and scattered cytoplasmic oil droplets are the main characteristics of mature head cells. The secretion which accumulates in the subcuticular space stains positively for hydrophilic and lipophilic substances, with lipids prevailing for fully peltate trichomes. Histochemistry and thin layer chromatography detected terpenes and alkaloids. Fehling's test to detect of sugars in the secretion was negative. CONCLUSIONS: The continuous presence and activity of peltate trichomes on the ovary of Z. montana from early budding through to flowering and fruiting set, and its main chemical components, alkaloids and terpenes, suggest that they serve a protective function and are not related to the floral nectar source or to improving nectar quality.