Pericarp Ontogeny Of Tapirira Guianensis Aubl. (anacardiaceae) Reveals A Secretory Endocarp In Young Stage

Most species of Anacardiaceae have drupes containing secretory structures.. The substances produced by these structures may have importance to industry and folk medicine, and may even cause allergenic effects. This work describes the ontogeny of pericarp of Tapirira guianensis with an emphasis on the secretory structures present at different stages of development. Ovary and fruits in various stages of development were collected, fixed and processed for studies using light and scanning electron microscopy according to conventional techniques. Histochemical tests were employed to identify the major metabolites present in the tissues. The fruit is a drupe formed by exocarp, mesocarp containing secretory ducts and idioblasts, and endocarp with some lignified layers. Fruit growth occurs through the division and elongation of cells. The secretory ducts produce mainly phenols and lipids and are active during all stages of development. The secreted substances protect the fruit against pathogens and predators. In ripe fruits the cells of the mesocarp accumulate starch. This study is the first report of the presence of a secretory endocarp in young fruits of a species of Anacardiaceae. The substances produced by the endocarp in early developmental stages may play an important role in seed dispersal and germination.31

Floral morpho-anatomy and reproductive ecology of Spondias macrocarpa Engl. (Anacardiaceae), a vulnerable neotropical andromonoecious tree

The genus Spondias has a wide variety of sexual systems, with complete monoecious to polygamodioecious species. Spondias macrocarpa (Anacardiaceae) is an endemic species from Brazil, previously described as hermaphroditic, and considered vulnerable due to massive deforestation. However, to our knowledge, no further details on the breeding system or morphological studies are available. Herewith, we examine the floral morpho-anatomy and breeding system of this species to better understand the sexual function of different floral morphologies and the role of pollinators in its reproduction. We used light and scanning electron microscopy to study the morphology and anatomy of the flowers. Additionally, we carried out experiments on pollen viability, stigmatic receptivity, and controlled pollinations. We found both bisexual and male flowers in the same inflorescence, with the staminate flowers presenting cryptic unisexuality. Investment in bisexual flowers was higher than in male ones, although more than 70% of the inflorescence was made up of male flowers. Most male flowers are present in the proximal region of the inflorescence, and there is no temporal separation between male and female functions in bisexual flowers. We show that S. macrocarpa is andromonoecious with cryptic-monoecy and a self-incompatibility system. Under natural conditions, S. macrocarpa is probably a pollinator generalist, with small bees being its main floral visitors. High levels of pollen robbing and low levels of crossing by floral visitors may result in the observed low fruit set. Further studies on the pre- and post-pollination barriers and on the reproductive ecology of extant natural populations should elucidate the reason for such a low fruit set

Osmophores and floral fragrance in Anacardium humile and Mangifera indica (Anacardiaceae): an overlooked secretory structure in Sapindales

Background and aims: 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. Methods: 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. Key results: 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 Anacardium humile and monoterpenes in Mangifera indica. Conclusions: 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

Structural studies on the reproductive organs of Tapirira guianensis Aubl. (Anacardiaceae)

Orientadores: Sandra Maria Carmello Guerreiro, Silvia Rodrigues MachadoDissertação (mestrado) - Universidade Estadual de Campinas, Instituto de BiologiaResumo: Tapirira guianensis Aubl. (Anacardiaceae) conhecida popularmente como pau-pombo é uma espécie importante para uso madeireiro, medicinal e pode ser empregada na recuperação de áreas degradadas e de matas ciliares. Diversas substâncias são produzidas em estruturas secretoras específicas, sendo a presença de canais e cavidades secretoras característica universal da família. Na literatura é relatada a presença de disco nectarífero intraestaminal nas flores de muitas Anacardiaceae. No entanto, estudos anatômicos e histoquímicos recentes não comprovaram a secreção de néctar por esta estrutura. Além disso, nestes trabalhos foi verificada a persistência do disco intraestaminal no fruto maduro de T. guianensis, bem como a presença de um endocarpo secretor em frutos não maduros. Aliado a estes fatores, a ontogenia de frutos e sementes também é um tema pouco explorado frente ao grande número de espécies da família. A anatomia dos frutos de Anacardiaceae tem-se mostrado de grande valor sistemático. Assim, o presente trabalho consistiu em verificar a estrutura e ultraestrutura do disco glandular de flores e frutos de T. guianensis, bem como examinar a natureza da secreção produzida e estudar a ontogenia do fruto e da semente. Com os resultados alcançados foi possível determinar que a glândula floral das flores hermafroditas de T. guianensis é uma glândula de secreção mista, produzindo néctar com predominância de sacarose, lipídios e substâncias fenólicas. Ultraestruturalmente esta glândula passa por mudanças profundas ao longo do desenvolvimento da flor, inicialmente rica em amiloplastos e posteriormente plastídios desprovidos de amido e mitocôndrias repletas de gotículas de óleo, indicando a mudança da secreção antes predominantemente hidrofílica para lipofílica. No que diz respeito à anatomia do fruto, observou-se que o mesmo possui características típicas da tribo Spondieae, porém algumas delas indicam uma condição menos derivada da espécie, como ausência de opérculo e endocarpo relativamente fino quando comparado com outras espécies da mesma tribo. O pericarpo possui grande quantidade de canais resiníferos na região do mesocarpo, epicarpo produzindo substâncias fenólicas e lipídios, e por meio de microscopia eletrônica de transmissão pôde-se observar o modo de secreção, as organelas envolvidas do processo, bem como a estrutura celular. A semente provém de um óvulo único, anátropo, bitegumentado, crassinucelado de funículo longo. A semente não possui camada mecânica, tem a exotesta e endotégmen ricos em substâncias fenólicas, cristais no mesotégmen, desenvolvimento de paquicalaza parcial, hipóstase fenólica e lipídica, permanência de funículo aderido ao envoltório da semente, formando um arilo vestigial. O embrião armazena grande quantidade de proteínas e amido, indicando uma condição de transição da espécie, pois em Anacardiaceae as reservas são constituídas principalmente por proteínas e lipídios. O fruto e semente apresentaram algumas características anatômicas possivelmente relacionadas à dispersão como, formação de aerênquima no mesocarpo interno e na mesotesta, acúmulo de amido no mesocarpo. O trabalho traz contribuições no estudo de glândulas florais em Anacardiaceae e na área de ontogenia de frutos e sementesAbstract: Tapirira guianensis Aubl. (Anacardiaceae) well-known as "pau-pombo" is an important species with uses as timber, in medicine and can be employed recovering degraded areas and riparian forests. Several substances are produced in specific secretory structures and the presence of cavities and ducts are universal features in the family. In literature is reported the presence of an intrastaminal receptacular nectary in the flowers of many Anacardiaceae, however anatomical and histochemical recent studies did not support the nectar secretion by this structure. Furthermore, in other studies were found the persistence of intrastaminal receptacular gland in T. guianensis ripe fruits and the presence of a secretory endocarp in unripe fruits. Allied these factors, the ontogeny of fruits and seeds is also a relatively unexplored subject, in contrast with the large number of species from this family. The fruit anatomy of Anacardiaceae has proved great systematic value. Thus, the present study verified T. guianensis structure and ultrastructure of flowers and fruits intrastaminal gland and examined the nature of substances produced as well the fruit and seed development. With the results obtained it was possible provide that the T. guianensis floral gland of hermaphrodite flowers is a mixed secretion gland, producing nectar with sucrose predominance, lipids and fenolic substances. Ultrastructurally, this gland undergoes deep changes during the development of the flower, originally rich in amyloplasts and subsequently chloroplasts devoids of starch and mitochondria replete with oil droplets, indicating the change in the secretion predominantly hydrophilic to lipophilic. With respect to the fruit anatomy, it was observed characteristics typical of the tribe Spodieae, but some of them indicate a condition less derived of this species, such as lack of operculum and endocarp relatively thin compared to other species of the same tribe. Pericarp has a large number of resiniferous ducts in mesocarp region, and epicarp produces phenolic substances and lipids. By transmission electron microscopy it was possible to observe the way of secretion, the organelles involved in process and the cell structure. Seed comes from a single ovule, anatropous, bitegmic, crassinucelate with longe funicle. Seed does not have mechanical layer, has exotest and endotegmen rich in phenolic substances, crystals in mesotegmen, development of partial pachycalaza, phenolic and lipid hypostasis, permanence of the funicle attached to the seed-coat, forming a vestigial aril. The embryo stores large amounts of protein and starch, indicating a transitional condition in the species, because Anacardiaceae reserves comprehend mainly protein and lipids. Fruit and seed showed anatomical characteristics possibly related to dispersion, like formation of aerenchyma in the inner mesocarp and mesotest, starch accumulations in mesocarp. The work brings contributions in the study of Anacardiaceae floral glands and in the area of fruit and seeds ontogenyMestradoBiologia VegetalMestra em Biologia Vegeta

Floral morphology and anatomy of Anacardiaceae with emphasis on the secretory structures

Orientadores: Sandra Maria Carmello Guerreiro, Leonardo GalettoTese (doutorado) - Universidade Estadual de Campinas, Instituto de BiologiaResumo: Anacardiaceae é uma das maiores famílias de Sapindales, representada por 82 gêneros e aproximadamente 800 espécies distribuídas principalmente em áreas tropicais. Estudos moleculares sustentam a monofilia da família, que encontra-se atualmente subdividida em duas grandes subfamílias, Spondioideae e Anacardioideae. Anacardioideae é monofilética, no entanto, a subfamília basal (Spondioideae) é polifilética e ainda necessita de mais estudos a fim de esclarecer melhor as relações filogenéticas do grupo. O estudo da notável variação morfológica e anatômica dos órgãos reprodutivos de Anacardiaceae pode fornecer o suporte necessário no esclarecimento de suas relações filogenéticas infrafamiliares. A ocorrência de estruturas secretoras em flores e frutos na família é amplamente discutida na literatura, podendo ocorrer, por exemplo, nectários, canais, coléteres e idioblastos, no entanto, são raros os trabalhos que discutem a composição química dos compostos produzidos e sua provável evolução na família. Nesta tese realizamos o estudo morfoanatômico de flores pertencentes às duas subfamílias de Anacardiaceae e discutimos o papel dos nectários florais e osmóforos para a sistemática e sucesso reprodutivo da família. Além disso também realizamos o estudo ontogenético de flores pertencentes à subfamília Spondioideae buscando esclarecer a evolução de flores unisexuais e do gineceu do tipo pseudomonômero em Anacardiaceae. Os resultados obtidos encontram-se divididos em três capítulos: (1) Diversidade estrutural e das secreções produzidas pelos nectários florais e suas implicações para a evolução de Anacardiaceae; (2) Osmóforos de Anacardium humile e Mangifera indica (Anacardiaceae), sua descoberta e relação com os polinizadores em flores nectaríferas de Sapindales; (3) Desenvolvimento floral em Tapirira guianensis e Spondias tuberosa, com particular referência à evolução de flores unisexuais crípticas e da pseudomonomeria em Spondioideae. As características estruturais dos nectários florais são conservativas dentro da família e a proporção dos açúcares presentes no néctar são de valor sistemático limitado. No entanto, todas as espécies estudadas apresentam néctar contendo lipídios, podendo esta ser uma sinapomorfia para Anacardiaceae. Uma nova estrututra secretora é descrita para a família, os osmóforos localizados na face adaxial de duas espécies pertencentes à subfamília Anacardioideae, A. humile e M. Indica. Os osmóforos são formados por uma única camada de células epidérmicas e a sua análise ultraestrutural revelou algumas diferenças significativas entre as duas espécies. Um link entre as diferenças encontradas e as fragrâncias produzidas é condizente, uma vez que o perfume floral é predominantemente composto por sesquiterpenos em A. humile e monoterpenos em M. indica. É ressaltada a importância de diferentes estratégias utilizadas pelos representantes de Sapindales na atração de polinizadores. O estudo de desenvolvimento floral demonstrou que Tapirira e Spondias possuem um padrão similar, com desenvolvimento simétrico dos carpelos, contrastando com representantes de Anacardioideae, em que o desenvolvimento do gineceu segue padrão assimétrico. Flores funcionalmente unisexuais são encontradas em membros da duas subfamílias e também em ambas subfamílias a pseudomonomeria é resultado da redução dos carpelos estéreis. Spondioideae difere de Anacardioideae pela ausência de uma zona simplicada entre os estiletes e a presença de uma zona sinascidiada expondo o ápice floral. A pseudomonomeria é considerada uma apomorfia para Anacardiaceae, especialmente para a subfamília Anacardioideae. Ainda discute-se questões relativas à variações merísticas do gineceu em AnacardiaceaeAbstract: Anacardiaceae are one of the largest families of Sapindales, represented by 82 genera and approximately 800 species distributed mainly in tropical areas. Molecular studies support the monophyly of the family, which is currently subdivided into two major subfamilies, Spondioideae and Anacardioideae. Anacardioideae is monophyletic, however, the basal subfamily (Spondioideae) is polyphyletic and still needs further studies in order to clarify its phylogenetic relationships. The study of the remarkable morphological and anatomical variation of the reproductive organs in Anacardiaceae may provide the necessary support to clarify its infrafamiliar phylogenetic relationships. The occurrence of secretory structures in flowers and fruits of the family is widely discussed in literature, for example, nectaries, ducts, coleters and idioblasts, however, there are just a few works discussing the chemical composition of their secretions and their evolution in the family. In this thesis we performed the morphoanatomic study of flowers belonging to the two subfamilies of Anacardiaceae and discussed the role of floral nectaries and osmophores for the reproductive success and systematic of the family. In addition, we also carried out the ontogenetic analysis of flowers belonging to the subfamily Spondioideae, seeking to clarify the evolution of unisexual flowers and the pseudomonomerous gynoecium in Anacardiaceae. The results obtained are grouped into three chapters: (1) Diversity of floral nectary secretion and structure, and implications for their evolution in Anacardiaceae; (2) Osmophores of Anacardium humile and Mangifera indica, their discover and relation to pollination in nectariferous flowers of Anacardiaceae; (3) Flower development in Tapirira guianensis and Spondias tuberosa, with particular reference to the evolution of the cryptic unisexual flowers and pseudomonomery in Spondioideae. The structural characteristics of floral nectaries are conservative within the family and the proportion of sugars of the nectar is of limited value for the systematic. However, all studied species have nectar containing lipids, which may be considered a synapomorphy for Anacardiaceae. A new secretory structure is described for the family, the osmophores located on the adaxial side of two species of the subfamily Anacardioideae, A. humile and M. indica. The osmophores are constituted by a single layer of epidermal cells and their ultrastructural analysis revealed some significant differences between the two species. A link between those differences and the produced fragrances is consistent, since floral scent is predominantly composed of sesquiterpenes in A. humile and monoterpenes in M. indica. It is emphasized the importance of different strategies used by the representatives of Sapindales in the attraction of pollinators. The floral development study showed that Tapirira and Spondias have flowers with a similar ontogenetic pattern, with symmetrical development of the gynoecium, contrasting with representatives of Anacardioideae, in which the development follows an asymmetric pattern. Functionally unisexual flowers are found in members of both subfamilies and the pseudomomery is a result of the reduction of sterile carpels. Spondioideae differs from Anacardioideae by the absence of a symplicate zone between the styles and the presence of a sinascidiate zone exposing the floral apex. Pseudomonomery is considered an apomorphy for Anacardiaceae, especially for the subfamily Anacardioideae. Issues related to meristic variations of the gynoecium in Anacardiaceae are also discussedDoutoradoBiologia VegetalDoutora em Biologia Vegetal140358/2017-0CAPESCNP

Secretory Patterns in Colleters of Apocynaceae

Colleters of Apocynaceae are glands related to different types of protection of vegetative and floral meristems through the production of mucilage or a mixture of many different compounds. Although several anatomical papers have shown histological and histochemical aspects of colleters of the family, almost nothing is known about their secretory process. In this study, we analyzed two types of colleters in Apocynaceae: one produces mucilage and lipophilic compounds, while the other produces an exclusively mucilaginous secretion. The secretory epidermis of the colleters of Allamanda schottii and Blepharodon bicuspidatum has a dense cytoplasm with organelles responsible for the production of mucilage and lipids. This heterogeneous secretion is released through granulocrine and eccrine mechanisms and is temporarily stored in a subcuticular space before crossing the cuticle. Conversely, colleters of Mandevilla splendens and Peplonia axillaris produce only mucilage and have a very different secretory apparatus. The mechanism of secretion is granulocrine, and the exudate is firstly accumulated in a large periplasmic space and later in an intramural space before crossing the cuticle. Notably, the structure of the cuticle varies according to the secretion composition. Although the colleters of the family are histologically similar, this study demonstrates a metabolic and subcellular variability previously unknown for Apocynaceae

New perspectives on secretory structures in Clusia (Clusiaceae - Clusiod clade): production of latex or resins?

The Clusioid clade is quite diverse, encompassing the five families Clusiaceae, Bonnetiaceae, Calophyllaceae, Hypericaceae, and Podostemaceae. However, its members have important synapomorphies, such as the presence of xanthones, polyisoprenylated benzophenones, and quinones, as well as schizogenous secretory ducts and cavities, and tenuinucellate ovules. Owing to the milky appearance of the exudate present in ducts of these families (except Bonnetiaceae), many authors have referred to this secretion as latex, while others have preferred to use the term resin. Faced with this confusion about the description of the exudate present in Clusiaceae, the aim of this study was to identify which classes of substances are being produced by those structures in the genus Clusia, compared with other members of Clusiaceae and the Clusioid clade in general. Furthermore, we describe the structure and distribution of the secretory ducts in species of Clusia, as well as a detailed histochemical study. These secretory ducts secrete an exudate of variable composition, but predominantly terpenic. We conclude that Clusia, as well as some representatives of the Clusioid clade, do not have latex but resiniferous secretory ducts.The accepted manuscript in pdf format is listed with the files at the bottom of this page. The presentation of the authors' names and (or) special characters in the title of the manuscript may differ slightly between what is listed on this page and what is listed in the pdf file of the accepted manuscript; that in the pdf file of the accepted manuscript is what was submitted by the author

Presumed domatia are actually extrafloral nectaries on leaves of Anacardium humile (Anacardiaceae)

Presumed domatia are actually extrafloral nectaries on leaves ofAnacardium humile (Anacardiaceae), a species from the Brazilian savannah, is associated with foraging ants, which leads to the description of the structure of its leaves as domatia. However, the ants collect secretions on the leaves, a phenomenon that inspired us to elucidate this structure. To this end, we conducted an analysis of the distribution, micromorphology, anatomy and histochemistry of these supposed domatia. They are located in the axil of thicker leaf veins and are composed of glandular trichomes which exude glucose; they are, therefore, described here as extrafloral nectaries (EFNs). The nectariferous trichomes are located at the same level as the other epidermal cells on young leaves, while in mature leaves they were found in depressions of the leaf blade. The secretory activity of EFNs is limited to juvenile phases, becoming degenerated in mature leaves. Carbohydrates and lipophilic compounds were detected in the produced secretion. It can be concluded that the presence of ants foraging leaves to collect from EFNs indicates a mutualistic interaction, with the plant providing a sugar resource to the ants in return for protection against herbivory since they are present in developing leaves

Foliar colleters on Anacardiaceae: first report for The family

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)Colleters are secretory structures widely distributed in eudicots and with taxonomic value in many families. Although glandular trichomes have been described in some Anacardiaceae species, the chemical characterization of their secretions is scarce and to date there are no reports on colleters. Light microscopy and scanning electron microscopy were used to study the distribution and structure of colleters on the vegetative buds of Anacardium humile A. St.-Hil., Lithraea molleoides (Vell.) Engl., Spondias dulcis Parkinson, and Tapirira guianensis Aubl., and to characterize their secretory products histochemically. In all of these Anacardiaceae species, colleters are multicellular and multiseriate ovoid or club-shaped glandular trichomes of protodermic origin, present on both surfaces of leaf primordia. They reach the secretory phase at early stages of leaf development, after which they gradually degenerate, become brown, and fall off. Histochemical tests indicate that the secretion within the glandular cells and outside the trichomes is a complex mixture containing mucilage, fatty acids, and phenolic compounds, which are secretory products that can play an important role in the protection of meristems against desiccation and attack by pathogens. Therefore, the distribution of these glandular trichomes, their short-life, the chemical nature of their secretions and their presumed functions support their being classified as colleters.Colleters are secretory structures widely distributed in eudicots and with taxonomic value in many families. Although glandular trichomes have been described in some Anacardiaceae species, the chemical characterization of their secretions is scarce and to945337346FAPESP - FUNDAÇÃO DE AMPARO À PESQUISA DO ESTADO DE SÃO PAULOCAPES - COORDENAÇÃO DE APERFEIÇOAMENTO DE PESSOAL DE NÍVEL SUPERIORFundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)03/13556-514/18002-200/12469-3SEM INFORMAÇÃOWe are grateful to Prof. Jorge Yoshio Tamashiro (Universidade Estadual de Campinas, Brazil) for plant identification. We also thank the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) for their financial support (FAPESP 03/13556-5, 14/18002