Abstract in English:ABSTRACT 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.
Abstract in English:ABSTRACT Lamiaceae contains many species known for their aromatic properties that are produced by the production of essential oils in glandular trichomes. Hyptis is one of the most common genera of Lamiaceae in the Brazilian flora, and includes several species with medicinal value. However, studies on the morphology and functioning of their glandular trichomes are lacking. We analyzed the morphology, histochemistry and ultrastructure of the glandular trichomes in leaves of H. villosa, emphasizing the differential distribution of actin filaments and microtubules in cells secreting hydrophilic and lipophilic compounds. Four morphotypes of glandular trichomes were identified. Total lipid, terpenes, alkaloids, phenolic compounds, proteins and polysaccharides were histochemically detected in all morphotyes. This evidences the mixed nature of the secretions of this species, although there are differences in the prevalence of lipophilic and hydrophilic components among the glandular morphotypes and among the cells of the same trichome. The actin microfilaments are more abundant in cells that secrete mainly hydrophilic compounds, and microtubules predominate in cells that secrete lipophilic compounds. Our results corroborate the correlation between the glandular morphotype and the composition of the secretion produced, with a differential distribution of the cytoskeletal elements according to the prevalence of either hydrophilic or lipophilic substances.
Abstract in English:ABSTRACT The genus Zeyheria (Bignoniaceae) comprises only two species, both of which have been described as possessing a reduced and non-functional nectary disk. Despite the importance of this evolutionary change in the floral nectary, these functional assumptions have been based on disk size and on the distribution, abundance and histochemistry of corolla-borne trichomes. By combining methods on light and electron microscopy, here we investigated the functionality of the reduced nectary disk and describe all of the tissues and structures of the nectar chamber in order to determine the sites of floral nectar secretion in both Zeyheria species. . Our data find the floral nectary traits of both species to be very similar, although differing in their cellular contents. Subcellular evidence in both species indicated that disk, stipe and petal axils were, predominantly, involved in hydrophilic secretion, while capitate glandular trichomes produced lipophilic secretion and papillae produced mixed secretion. Our study shows that in spite of its reduced size, the reduced disk functions in nectar secretion in both species of Zeyheria. This kind of nectary system is a novelty for Bignoniaceae, since it comprises several tissues and structures functioning in an integrated fashion.
Abstract in English:ABSTRACT Floral secretory structures are usually associated with the attraction of pollinators, but may also play an important role in the mechanisms of plant protection. This study aimed to show the diversity of secretory structures present in the developing flowers of 15 legume species belonging to different clades and to associate them with functions other than the pollinator attraction. Buds, flowers and developing axis of inflorescence were processed for surface, histological, and ultrastructural analyses. The species investigated displayed a wide diversity of secretory structures in developing flowers such as phenolic cells and/or tissues, mucilaginous cells, secretory cavities, secretory trichomes and colleters. Each type of secretory structure exhibited variation in morphology and location in the flower and/or axis of inflorescence depending on the species. Special mucilage cells, secretory cavities, secretory trichomes and colleters have great potential for comparative morphological studies due to their diversity of forms or restricted occurrence to certain taxa, contributing to a more robust morphological data base for the new clades emerging in Leguminosae. The scarcity of reports about floral secretory structures of Leguminosae seems to be more related to deficient sampling than to the absence of such structures in the group, which highlights the need for further investigation.
Abstract in English:ABSTRACT Dipteryx, Pterodon and Taralea are legume genera known for secreting oils, produced in secretory canals and cavities, with medicinal properties. We analyzed the distribution, morphology and histochemistry of these glands in leaves and stems of Dipteryx alata, Pterodon pubescens and Taralea oppositifolia, three Neotropical species, using standard techniques for anatomy and histochemistry. Secretory spaces, i.e. secretory cavities and canals, exhibited a wide lumen and a single layer of epithelium. Digitiform epithelial cells, forming trabeculae protruding into the lumen, were seen in all three species. Secretory cavities with a rounded or oval lumen and secretory canals with an elongated lumen in longitudinal sections were found only in T. oppositifolia. In D. alata and P. pubescens, only secretory cavities were found. In P. pubescens, secretory cavities occurred in the leaf blade margin. In T. oppositifolia, secretory spaces were much more numerous than in the other two species. Terpenes, total lipids, phenolic compounds, alkaloids and polysaccharides were detected in the secretory spaces of the three species. The abundance of secretory spaces, the presence of canals in T. oppositifolia and the position of cavities in P. pubescens are features with potential diagnostic value for their respective genera.
Abstract in English:ABSTRACT The genus Chamaecrista contains ca. 330 species organized into six sections, of which section Chamaecrista is the second largest (ca.75 species) distributed among six series, while the section Caliciopsis contains only two species. Colleters have been described in the genus Chamaecrista and they show potential taxonomic and phylogenetic significance. They are associated with lubrication, desiccation prevention and protection from microbial attacks of young developing organs. Although six types of colleters have been described for the genus Chamaecrista, there have been no studies focusing on the diversity of colleters in the sections Chamaecrista and Caliciopsis. Samples from developing leaves and flowers of both sections were obtained from herbarium and field collections and subjected to standard methodologies for both light and scanning electron microscopy. Histochemical tests were also performed to determine the nature of the exudates. Five types of non-vascularized colleters were found: short digitiform, long digitiform, club-shaped, pyriform and short bottle-shaped. Polysaccharides, pectins, lipids and proteins were detected in the exudates of all types of colleters. Among the five types of colleters observed, pyriform is a novelty for Chamaecrista, reinforcing the significant morphological diversity of these secretory structures in this genus.
Abstract in English:ABSTRACT This study aimed to obtain new information about the distribution, morphology and content of secretory trichomes present in the vegetative and reproductive organs of four species of Moraceae: Artocarpus heterophyllus, Dorstenia cayapia, Maclura tinctoria and Sorocea bonplandii. Stem, leaf, flower and inflorescence samples were processed for scanning electron and light microscopy. The species have secretory trichomes on the leaf blade, petiole and stem and also on the inflorescence and flowers of D. cayapia and S. bonplandii and on the peduncle of the inflorescence in A. heterophyllus. These trichomes are of the capitate type in all species, but A. heterophyllus also possess peltate secretory trichomes. Both trichome types exhibit a multicellular head and a unicellular peduncle. Peduncle size and head cell number varies according to species. Peltate trichomes located in depressions were found only in species of Artocarpus, and so should be of diagnostic value for the genus. In all species the trichomes secrete polysaccharides, alkaloids and lipids; phenolic compounds occur in A. heterophyllus and M. tinctoria trichomes. These trichomes probably act in the protection against herbivores, pathogens, desiccation and/ or UV radiation. Moreover, the trichomes found in the inflorescence of D. cayapia may exert a function in pollen grain adhesion.
Abstract in English:ABSTRACT Extrafloral nectaries (EFNs) are specialized structures that produce and release nectar and are located on leaves, cotyledons and, more seldomly, stems. Peculiar leaf nectaries have been described for thirty-three Neotropical species of Gentianaceae, while stem nectaries have been reported for only nine. The aim of this study was to verify the occurrence of stem EFNs within Gentianaceae and investigate the existence of a correlation between their occurrence and the geographical distribution of species. Samples of internodal regions from field and herbarium specimens were submitted to standard light microscopy techniques. Data regarding the geographical distribution of species were acquired from herbarium specimens and the literature. A total of 37 species were investigated, representing 25 genera distributed among five tribes. Nectaries, composed of modified epidermal cells, were observed in 16 species restricted to the Neotropical tribes Helieae, Saccifolieae, Potalieae and Coutoubeinae; exceptions were Cicendia quadrangularis and Zygostigma australe, which both occur in the Neotropics but do not possess EFNs. These results demonstrate that stem EFNs are common among Neotropical taxa of Gentianaceae, and are typically absent from taxa in temperate regions.
Abstract in English:ABSTRACT Secretory structures are formed by many types of plants and are present on numerous different organs. Among the many types of known secretory structures, colleters are predominant on plants of the Rubiaceae. One remarkable characteristic of secretory tissues is the export of exudates; however, the precise mechanism involved in this process is still unclear. To better understand the mechanisms of exudate externalization it is necessary to understand the ultrastructure and dynamics of the outer cell wall of the secretory structures during the secretory process, and so we investigated these aspects of the colleters of Bathysa nicholsonii. The outer cell wall (OCW) exhibits multiple layers: a basal polysaccharide rich layer; a cuticular membrane, which is subdivided into arborescent and reticulated layers; and a thin cuticle proper. The structural organization of the OCW is changed during secretion passage, which is mainly related to the development of a secretion accumulation site on the polysaccharide rich layer. Secretion dynamics is driven by the organization and disruption of the secretion accumulation site. The results show that the OCW of the colleters of B. nicholsonii is a dynamic structure with an active role in secretion externalization via constant structural reorganization directly related to secretion passage.
Abstract in English:ABSTRACT Flowers of Asclepiadoideae are notable for possessing numerous nectaries and elaborate coronas, where nectar can accumulate but is not necessarily produced. Given the complexity and importance of these structures for reproduction, this study aimed to analyze the ontogeny of the corona, the structure and position of nectaries and the histochemistry of the nectar of species of Asclepiadeae. Two types of coronas were observed: androecial [C(is)] and corolline (Ca). The development of the C(is)-type of corona initiates opposite the stamens in all species examined with the exception of Matelea in which it begins to develop as a ring around the filament tube. Despite their morphological variation, coronas typically originate from the androecium. A notable difference among the studied species was the location of the nectaries. Primarily, they are located in the stigmatic chamber, where nectar composed of carbohydrates and lipids is produced. A secondary location of nectaries found in species of Peplonia and Matelea is within the corona, where nectar is produced and stored, composed of carbohydrates and lipids in Peplonia and only carbohydrates in Matelea. The functional role of nectar is related to the location of its production since it is a resource for pollinators and inducers of pollen germination.
Abstract in English:ABSTRACT Floral nectaries are linked with animal pollinators and floral specialization, characterizing some plant groups, such as the mostly Neotropical Vochysiaceae. The phylogeny of these plants has been revised based on molecular data, but their conspicuous spurs were mostly neglected. The appendicular or receptacular origin of these spurs has been discussed but without ontogenetic or anatomical study, and the association between spur structure and pollination systems was never assessed. We studied the spurs of species of four genera of Cerrado Vochysiaceae (tribe Vochysieae), documenting their vascularization and comparing their structure to previously described pollination systems. The spur structure was roughly similar among species, and vascularization confirmed their appendicular origin. Subepidermic secretory tissue surrounded the lumen of the spur and nectar was secreted through nectarostomata. Besides differences in size and cuticle striation, no other association between spur structure and pollination system was observed. However, the structure is somewhat distinct in Qualea parviflora, which sports a pluriseriate internal epidermis padding the spur lumen, which may provide protection against damage by the large bees that visit these relatively small flowers. The conservative histology and vasculature somewhat confirmed the monophyly of the tribe Vochysieae and the appendicular origin of the spur in the Cerrado Vochysiaceae.
Abstract in English:ABSTRACT Nectaries are specialized structures that secrete nectar. Several species of Chamaecrista possess nectaries on the petiole, which have been shown to vary widely in morphology and the chemical nature of their secretion. However, a comprehensive investigation of the nectaries of the clade formed by sect. Chamaecrista, Caliciopsis and Xerocalyx has yet to be performed. Our study aimed to confirm whether or not the leaf glands of species of this clade are in fact nectaries, determine the chemical nature of their secretion and expand the morphoanatomical database on leaf nectaries in Chamaecrista with the intention of contributing to the taxonomy and phylogeny of the genus. Samples from herbarium and field-collected material were subjected to standard methods for light and scanning electron microscopy. Four different forms of nectaries were observed: urceolate, patelliform, verruciform and cupuliform. The nectaries were found to comprise a single-layered epidermis, nectary parenchyma, subnectary parenchyma and vascularization. Polysaccharides, lipids, phenolic compounds and proteins were detected in secretions. Although anatomical similarities were observed among the studied species, their morphology differed. Moreover, the glands are indeed nectaries and are similar to those observed in other species of the genus Chamaecrista. These data hold potential taxonomic usefulness for the studied sections.
Abstract in English:ABSTRACT Leaf glands are common structures in Malpighiaceae and exhibit great morphological diversity, yet information on their anatomy, secretion and type of visitors remains scarce. The aim of this study was to describe the distribution, anatomical development and chemical and functional properties of leaf glands of Banisteriopsis muricata (Malpighiaceae). Leaves at different stages of development were collected and processed according to standard techniques for light and scanning electron microscopy. Secretion composition was determined by histochemical tests and test-strips, while gland funciton was determined by field observation of interactions with visitors. Leaf glands were located on the petiole and on the abaxial base of the leaf blade. The gland secretion was found to be a protein-rich nectar that was foraged upon by ants ( Solenopsis); it was found accumulated in subcuticular spaces without pores or stomata for its release. Leaf glands were found to develop from protoderm and ground meristem, and consisted of typical secretory epidermis, nectariferous parenchyma and vascularized subnectariferous parenchyma. Therefore, it can be concluded that the distribution, chemical nature of secretion and anatomy of leaf glands of B. muricata characterize them as EFNs, while foraging by ants indicate a mutualistic relationship that possibly protects the plant against herbivores.
Abstract in English:ABSTRACT This study aimed to anatomically characterize the extrafloral nectaries (EFNs) of annatto (Bixa orellana) and determine the composition of its nectar in order to better understand their structure and function during organ development. Standard light and scanning electron microscopy techniques were used for anatomical analysis, and test-strips and a refractometer for determining nectar composition. Both receptacle and stem EFNs were found to possess similar anatomy and nectar composition, and to secrete from early to advanced developmental stages of the organs to which they are associated. EFNs consisting of uniseriate epidermis, nectariferous parenchyma and subnectariferous parenchyma were found located where vascular tissue is immersed. Some layers of nectariferous parenchyma exhibited sclerification and cells with phenolic compounds or calcium oxalate druses were present. Nectar exuded by stomata was acidic, diluted and found to contain sugar. The anatomical and histochemical features of annatto EFNs ensure their integrity and nectar secretion function during the development of buds, flowers and fruits.
Abstract in English:ABSTRACT Species of Apocynaceae stand out among angiosperms in having very complex flowers, especially those of asclepiads, which belong to the most derived subfamily (Asclepiadoideae). These flowers are known to represent the highest degree of floral synorganization of the eudicots, and are comparable only to orchids. This morphological complexity may also be understood by observing their glands. Asclepiads have several protective and nuptial secretory structures. Their highly specific and specialized pollination systems are associated with the great diversity of glands found in their flowers. This review gathers data regarding all types of floral glands described for asclepiads and adds three new types (glandular trichome, secretory idioblast and obturator), for a total of 13 types of glands. Some of the species reported here may have dozens of glands of up to 11 types on a single flower, corresponding to the largest diversity of glands recorded to date for a single structure.
Abstract in English:ABSTRACT Elaiophores are floral glands that secrete non-volatile oils as a reward for their pollinators. Their secretions mediate a specialized interaction between oil-producing plants and a few species of oil-collecting bees - Apidae and Melittidae. The present review focuses on the morphological and functional aspects of these secretory structures. We also provide information on their chemistry and pollination ecology. Our survey is organized taxonomically among the plant families for which elaiophore occurrence has been confirmed -for the monocots, Iridaceae and Orchidaceae, and for the eudicots, Calceolariaceae, Cucurbitaceae, Krameriaceae, Malpighiaceae, Plantaginaceae, Primulaceae, Scrophulariaceae, Solanaceae and Stilbaceae. Most oil flowers are zygomorphic, bisexual, nectarless and present their elaiophores on the protective whorls or on the androecium. Trichomal elaiophores are reported in all of these families except Krameriaceae and Malpighiaceae, and they vary in the density and morphology of the trichomes. Epithelial elaiophores occur in some monocot representatives and in Krameriaceae and Malpighiaceae, and are similar among species. An intermediate type of elaiophore is reported in some species of Orchidaceae. Regardless of elaiophore type, these glands have similar subcellular features and secretion-release mechanisms. Finally, we note the dearth of information on elaiophore ultrastructure and functioning for several plant families.
Abstract in English:ABSTRACT In many glandular structures, departure from the cell is only one step in the process of exudate release to the plant surface. Here the set of events that lead nectar to the external environment is presented and discussed mainly for stomata-free nectaries. After being synthesized, the nectar or some of its component needs to be released to the environment where it performs its functions. Nectar precursors derived from cell metabolism need to cross several barriers, such as the cell membrane and cell wall, in order to become nectar. Then the nectar must cross the cuticle or pass through stomata in order to be offered to plant mutualists. Release through stomata is a simple mechanism, but the ways by which nectar crosses the cuticle is still controversial. Hydrophilic pathways in the cuticle and repetitive cycles of rupture or cuticle detachment are the main routes for nectar release in stomata-free nectaries. In addition to nectar, there are other exogenous secretions that must leave the protoplast and reach the plant surface to perform their function. The ways by which nectar is released discussed herein are likely relevant to understanding the release of other hydrophilic products of the secretory process of plants.