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MAIN CONCLUSION: New findings are presented for Chaerophyllum coloratum L. on the volatile composition of the essential oil, based on data of hydrosol and fresh plant material, light and electron microscopy of leaves, and cytotoxic and antiviral activity. The widespread Apiaceae family includes many well-known and economically important plants that are cultivated as food or spices. Many produce essential oils and are generally a source of secondary metabolites and compounds that have numerous applications in daily life. In this study, the chemical composition of volatile organic compounds (VOCs), ultrastructure and biological activity of the Mediterranean endemic species Cheaerophyllum coloratum L. are investigated, as literature data for this plant species are generally very scarce. The essential oil and hydrosol were extracted from the air-dried leaves by hydrodistillation and the chemical composition of both extracts was analysed by GC-MS in conjunction with headspace solid-phase microextraction (HS-SPME) of VOCs from the hydrosol and the fresh plant material. In the composition of the essential oil, the oxygenated sesquiterpenes spathulenol and caryophyllene oxide were the most abundant components. In the fresh plant material, non-oxygenated sesquiterpenes dominated, with ß-caryophyllene and germacrene D being the main components. The hydrosol was dominated by monoterpenes, with the oxygenated monoterpene p-cymen-8-ol being the most abundant. Light and electron micrographs of the leaf of C. coloratum show secretory structures, and we hypothesize that glandular leaf trichomes, secretory epidermal cells and secretory canals are involved in the production of volatiles and their secretion on the leaf surface. Since the biological potential of C. coloratum is poorly investigated, we tested its cytotoxic activity on cancer and healthy cell lines and its antiviral activity on plants infected with tobacco mosiac virus (TMV). Our results dealing with the composition, ultrastructure and biological activity show that C. coloratum represent a hidden valuable plant species with a potential for future research.

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Secretory structures in plants play a crucial role in producing bioactive compounds. Despite the potential of the Swartzia genus, comprehensive studies in this context are still scarce. Swartzia is a legume tree (Fabaceae) that occurs in the Brazilian Atlantic Forest, a biodiversity hotspot, and includes species such as Swartzia flaemingii. Therefore, we aim to achieve: (1) identify and characterize the key secretory sites responsible for saponin production in S. flaemingii leaflets; (2) confirm the presence of saponins in S. flaemingii leaves by comparing them with known chemical profiles of other Swartzia species; (3) assess the potential hemolytic and cytotoxic effects of crude leaf extracts. Our investigation unveils the presence of phenolic idioblasts, mucilage cells, and articulate laticifers, which play pivotal roles in defense and adaptation. Notably, we report the first-ever ultrastructural details of laticifers in a legume species. Additionally, oleanane-type saponins were identified in the leaves, giving insights into the chemotaxonomic profile of Swartzia. The crude extracts show low cytotoxicity levels, showcasing as a promising alternative source of saponins. This investigation reinforces the importance of conserving plants in threatened regions like the Atlantic Forest, a global biodiversity hotspot facing substantial anthropogenic pressures.

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Santolina impressa is an aromatic Asteraceae species endemic to Portugal, traditionally used for its anti-inflammatory properties. The aim of this study was to characterize S. impressa secretory structures, analyze the essential oil (EO) from the aerial organs, and evaluate its antiviral activity against herpes simplex viruses HSV-1 and HSV-2. Secretory structures were investigated by light and scanning microscopy, and the secretion was histochemically characterized. The EO from the aerial organs in full blooming was analyzed by gas chromatography with flame ionization detection and gas chromatography-mass spectrometry. Antiviral assays were performed by direct contact with viral suspensions (virucidal effect), and in infected Vero E6 cells, at different time periods during the viral replication cycle. Two types of secretory structures were described, biseriate glandular trichomes and secretory ducts, producing an oleoresin and a resin rich in flavonoids, respectively. Fifty compounds were identified in S. impressa EO, accounting for 87% of the total constituents. Monoterpenes constituted the main EO fraction (82%), with ß-pinene (13%) and ß-phellandrene (10%) being their major components. The EO interacted with HSV-1 and HSV-2 in a dose-dependent manner, thereby inactivating both viral infections. The EO did not evidence a virucidal effect but inhibited the HSV-1 and HSV-2 infection in Vero cells in a dose-dependent manner. However, further studies are needed to investigate the mode of action in the replication cycle.

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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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Croton gratissimus (Lavender croton) possesses three distinct secretory structures. These include lepidote and glandular trichomes and non-articulated unbranched laticifers. The lepidote trichomes form a dense indumentum on the abaxial surface of the leaves and canopy the glandular trichomes. Although assumed to be non-glandular, transmission electron microscopy (TEM) indicated high metabolic activity within the stalk and radial cells. Glandular trichomes are embedded in the epidermal layer and consist of a single cell which forms a prominent stalk and dilated head. Laticifers occur on the mid-vein of leaves and are predominantly associated with vascular tissue. In the stems, laticifers are associated with the phloem and pith. Both trichome types and laticifers stained positive for alkaloids, phenolic compounds, and lipids. Positive staining for these compounds in lepidote trichomes suggests their involvement in the production and accumulation of secondary metabolites. These metabolites could provide chemical defense for the plant and potentially be useful for traditional medicine.

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Histochemical analysis is essential for the study of plant secretory structures whose classification is based, at least partially, on the composition of their secretion. As each gland may produce one or more types of substances, a correct analysis of its secretion should be done using various histochemical tests to detect metabolites of different chemical classes. Here I describe some of the most used methods to detect carbohydrates, proteins, lipids, phenolic compounds, and alkaloids in the secretory structures.

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Coryanthes is one of the most fascinating genera of Stanhopeinae (Orchidaceae) because of its complex pollination mechanism and the peculiar structures of its flowers. Although Coryanthes macrantha is widely studied, investigation of the secretory structures and floral biology is important to understand the mechanisms and ecology of pollination, which deserve attention despite the difficulties of collecting fertile material in nature. We conducted a morpho-anatomical analysis of the floral and extrafloral secretory structures of C. macrantha to better understand the secretory structures, contribute to the knowledge of its floral biology and/or pollination processes and understand the ecological function of these structures. The analysis revealed that C. macrantha has epidermal osmophores with unicellular papillae that were foraged by male Eulaema bees, floral nectaries in the sepals and extrafloral nectaries in the bracts. In both the floral and extrafloral nectaries, the nectar is exuded by the stomata. Azteca ants foraged the bract and sepal nectaries in pre-anthesis and post-anthesis. We also described the secretory epidermis of pleuridia, and the mode of secretion of osmophores and nectaries and found that they attract specific foraging agents.

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Rhaponticum carthamoides (Willd.) Iljin is not only a source of phytosteroids and flavonoids, but is also source of essential oil (EO). This study evaluated the volatile metabolic constituents and histochemical localization of root and rhizome essential oils (EOs) from R. carthamoides populations wild-grown in Russia and cultivated in Bulgaria. The performed histochemical analysis confirmed the presence of lipophilic substances in the secretory ducts of the examined roots and rhizomes. Both EOs were obtained by hydrodistillation and further analyzed by gas chromatography with mass detection. The results showed differences between the chemical compositions of both EOs. Forty-six components were tentatively identified in R. carthamoides oil from the wild population, with ß-selinene (4.77%), estragole (6.32%), D-carvone (6.37%), cyperene (8.78%), and ledene oxide (11.52%) being the major constituents. In the EO isolated from cultivated R. carthamoides, twenty-three compounds were tentatively identified, with humulene (7.68%), ß-elemene (10.76%), humulene-1,2-epoxide (11.55%), ledene oxide (13.50%), and δ-elemene (19.08%) predominating. This is the first report describing the histolocalization and chemical profile of EO from R. carthamoides cultivated in Bulgaria. Further research on the cultivation of R. carthamoides in Bulgaria would affect the relationship between its chemical composition and pharmacological effects.

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Nectar plays important roles in the relationship between plants and other organisms, both within pollination systems and as a defense mechanism. In the latter case, extranuptial nectaries (ENNs) usually attract patrolling arthropods that reduce herbivory. ENNs have been frequently reported within the "xeric clade" of Bromeliaceae, but their occurrence in other groups of bromeliads is largely unexplored, especially considering their position, secretory activity and structure. After observing the presence of ants constantly patrolling the inflorescences of Pitcairnia burchellii Mez, we searched for the presence, secretory activity, and structure of ENNs in this species. We also provide a brief review of the occurrence ENNs in Bromeliaceae. The distribution of nectaries was assessed using ant-exclusion experiments, while structural analysis was performed using standard methods for light and scanning electron microscopy. The presence of sugars in the secretion was assessed by thin-layer chromatography and glucose strip tests. Nectaries in P. burchelli are non-structured glands on the adaxial surface of floral bracts and sepals. Bracts and sepals are distinct spatial units that act over time in the same strategy of floral bud protection. Literature data reveals that ENNs might be more common within Bromeliaceae than previously considered, comprising a homoplastic feature in the family. Future perspectives and evolutionary and taxonomic implications are discussed.

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Colleters are secretory structures involved in the protection of young and developing plant organs. Although the presence of colleters in Gentianales is described as a synapomorphy, studies on the morphofunctionality of colleters and the mechanisms underlying the synthesis and release of colleter secretion in Gentianaceae are scarce. Here, we described the ontogeny and the morphological and functional aspects of colleters of Prepusa montana, revealed the nature of the key compounds present in the secretion, and explored the cellular aspects of the synthesis and release of secretion and senescence of colleters. Samples of the stem apical meristem with leaf primordium and young leaves; adult and senescent leaves were observed using light and electron microscopy. The colleters, located in the axil of the leaf, have a protodermal origin and develop asynchronously. They are digitiform, possessing a short peduncle and a secretory head containing homogeneous cells with dense cytoplasm and abundant endoplasmic reticulum and Golgi bodies. The secretion, composed of polysaccharides and proteins, is accumulated in schizogenous spaces and released through the separation of peripheral secretory cells and loosening of the external periclinal wall. Presumably, senescence is caused by programmed cell death. The morphoanatomical characterization of P. montana leaf colleters described here is the first record for the genus and the peculiar accumulation of colleter secretion in schizogenous spaces expanding our knowledge on the diversity of these secretory structures. Our results also provide insights into programmed cell death as an eminent topic related to secretory structures.

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Floral secretory structures have been reported for Gentianaceae; however, morphoanatomical studies of these glands are rare. We described the development and secretory activity of the colleters and nectaries throughout the floral development of Chelonanthus viridiflorus. We collected flower buds, flowers at anthesis, and fruits to be investigated using light and scanning electron microscopy. We performed histochemical tests on the secretion of colleters and used glycophyte to confirm the presence of glucose in nectar. Colleters are located on the ventral surface of sepals and nectaries occur in four regions: (i) the dorsal and (ii) ventral surfaces of sepals; (iii) apex of petals; and (iv) base of ovary. The colleters have a short peduncle and a secretory portion with homogeneous cells. They are active in flower buds and secrete polysaccharides and proteins. In flowers at anthesis, they begin to senescence presenting protoplast retraction, cell collapse, and lignification; these characteristics are intensified in fruit. The nectaries of sepals and petals have two to five cells surrounding a central cell through which the secretion is released. Nectaries are numerous, forming a nectariferous area on the dorsal surface of sepals, like that observed on petals, and can form isolated units on the ventral surface of sepals. They are active from flower buds to fruits. A region with secretory activity was identified at the base of the ovary. The secretion of colleters acts in the protection of developing organs, while nectaries are related to defenses against herbivores and the supply of nectar to potential robbers or pollinators.

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Croton L. (Euphorbiaceae) is the second largest genus of the Euphorbiaceae s.l. family, which is composed of about 1,200 taxa. Several species are widely used medicinally in various parts of the world to treat cancer, digestive problems, skin diseases, malaria, ulcers, and obesity. The leaves and stems are sold as dehydrated fragments, or in powder form, and used in the preparation of teas and various types of formulations. The aim of this study was to accurately characterize the leaf anatomy of Croton bonplandianus Baill. and Croton gracilipes Baill. and to perform histochemical analysis of laticifers to provide subsidies for botanical certification. To accomplish this, leaf blade and petiole samples were fixed and prepared following standard techniques for scanning electron and light microscopy. Histochemical tests were carried out on sections to detect the compounds present in the secretions. Results showed neutral and acidic lipids, as well as phenolic compounds, in the latex. In addition, leaf anatomical structures of these two species were studied for specific identification of morphological characters and for quality assessment of these medicinal plants.

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Orobanche picridis is an obligate root parasite devoid of chlorophyll in aboveground organs, which infects various Picris species. Given the high level of phenotypic variability of the species, the considerable limitation of the number of taxonomically relevant traits (mainly in terms of generative elements), and the low morphological variation between species, Orobanche is regarded as one of the taxonomically most problematic genera. This study aimed to analyse the taxonomic traits of O. picridis flowers with the use of stereoscopic and bright-field microscopy as well as fluorescence, scanning, and transmission electron microscopy. The micromorphology of sepals, petals, stamens, and pistils was described. For the first time, the anatomy of parasitic Orobanche nectaries and the ultrastructure of nectaries and glandular trichomes were presented. Special attention was paid to the distribution and types of glandular and non-glandular trichomes as well as the types of metabolites contained in these structures. It was demonstrated that the nectary gland was located at the base of the gynoecium and nectar was secreted through modified nectarostomata. The secretory parenchyma cells contained nuclei, large amyloplasts with starch granules, mitochondria, and high content of endoplasmic reticulum profiles. Nectar was transported via symplastic and apoplastic routes. The results of histochemical assays and fluorescence tests revealed the presence of four groups of metabolites, i.e. polyphenols (tannins, flavonoids), lipids (acidic and neutral lipids, essential oil, sesquiterpenes, steroids), polysaccharides (acidic and neutral polysaccharides), and alkaloids, in the trichomes located on perianth elements and stamens.

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Centaurea cyanus L. is a valuable source of many different bioactive substances. It is used in herbal medicine, but the structure of its organs used as raw material and secretory tissues has been insufficiently examined. The aim of this paper was to investigate the microstructure of C. cyanus flowers, bracts, stems and leaves with particular emphasis on secretory structures. Moreover, the main classes of secondary metabolites present in the secretion were identified and the taxonomic value of some micromorphological and anatomical features was analysed. Histochemical, micromorphological and ultrastructural analyses of aboveground organs of C. cyanus were carried out using light, fluorescence, scanning and transmission electron microscopy. The analyses revealed the presence of petal papillae and a characteristic cuticular pattern on the petals, stamens and stylar hairs. There were four types of non-glandular trichomes on the bracts, leaves and stem surfaces. The epidermal cells of the bracts contained prismatic calcium oxalate crystals. Two kinds of secretory structures, i.e. glandular trichomes and ducts, were observed in the C. cyanus organs. The glandular trichomes were located on the bract and stem surfaces, and the ducts were detected in the leaves and stems. Ultrastructural analyses of the epithelium of the ducts showed the presence of strongly osmiophilic insoluble phenolic material in vacuoles as well as moderately osmiophilic insoluble lipidic material in elaioplasts and vesicles. The results of histochemical assays showed a heterogeneous nature of the duct secretion, which contained essential oil, lipids, flavonoids, tannins and terpenes containing steroids.

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PREMISE: Laticifers have evolved multiple times in angiosperms and have been interpreted as a key innovation involved in plant defense mechanisms. In Malpighiaceae, laticifers were previously known from a single lineage of trees and shrubs, the Galphimia clade, but with detailed anatomical analyses here, we show that their distribution is broader in the family, also encompassing large clades of lianas. METHODS: From 15 genera, 70 species of Malpighiaceae were surveyed through careful anatomical ontogenetic analysis of roots, stems, and leaves and detailed histochemical tests to elucidate the nature of laticifers and latex in the family. RESULTS: Articulated anastomosing laticifers were encountered in roots, stems, and leaves of two distantly related megadiverse genera of Malpighiaceae lianas: Stigmaphyllon (stigmaphylloid clade) and Tetrapterys s.s. (tetrapteroid clade). From the apex downward, in Stigmaphyllon the laticifers are derived from the procambium and from the cambium during its early activity and are present in the outermost part of the vascular cylinder of stems and leaves and in the pericycle of roots, whereas in Tetrapterys s.s. they are derived from the ground meristem, procambium, and cambium throughout the plant body and are present in the cortex and pith, either the pericycle in roots or the outermost part of the vascular system in stems and leaves, and the primary and secondary phloem. CONCLUSIONS: Laticifers seem to have evolved at least three times independently in Malpighiaceae, once in a lineage of trees and shrubs and twice in two distantly related megadiverse lianescent lineages. Laticifer evolution in Malpighiaceae is homoplastic and may be related to increases in species diversification.

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Abstract Dipteryx odorata (Aubl.) Willd. and Taralea oppositifolia Aubl., Fabaceae: Dipterygeae, are two Amazonian species of great economic and pharmacological potential. The anatomy of these species, however, remains poorly studied. The aim of this work was to inventory leaf anatomical characteristics of D. odorata and T. oppositifolia and to locate and identify secretory structures and determine the main classes of metabolites they store. Vegetative branches were collected in Parque Ecológico de Gunma, Belém, state of Pará, Brazil. Some of the branches were destined for herborization while the remainder was submitted to standard protocols for anatomical analysis and histochemical tests. Both species were found to possess an unstratified epidermis, with D. odorata being amphistomatic and T. oppositifolia being hypostomatic, and dorsiventral mesophyll with spongy parenchyma and wide cellular space. The two species were also found to possess idioblasts and secretory cavities that produce a heterogeneous exudate consisting of polysaccharides, lipids, alkaloids and phenolic compounds. The species presented differences in leaf anatomy and chemical composition of the secretory structures, which may be useful in their differentiation.

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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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ABSTRACT: Medicinal plants are generally commercialized dried. However, temperatures used in their drying processes may interfere with the content and chemical composition of their essential oils. The Lippia origanoides possesses thymol in the essential oil that is stored in glandular trichomes. Thymol is a major component of economic importance in the chemical and pharmaceutical industry. The objective of this study was to evaluate the effect of the drying temperatures of L. origanoides leaves regarding trichome integrity, content, and chemical composition of the essential oil. The experimental design was completely randomized with four treatments defined by oven drying temperatures (40, 50, 60, and 70ºC) and four replications. Essential oil was extracted by hydro distillation and the essential oil contents were determined (%). Chemical composition of the oil was analyzed by gas chromatography coupled to mass spectrometry. Scanning electron microscopy was carried out to determine trichome integrity. Drying the L. origanoides leaves in a forced ventilation oven at 40°C minimized the loss of extracted essential oil content (17.5g kg-1) and relative thymol percentage. Leaves dried at the temperatures of 60 and 70ºC exhibited a higher percentage of ruptured trichomes and reduced essential oil content to 13.7g kg-1 and 11.8g kg-1, respectively.

RESUMO: As plantas medicinais são geralmente comercializadas secas, no entanto, as temperaturas utilizadas nos processos de secagem podem interferir no teor e composição química dos óleos essenciais destas plantas. A Lippia origanoides possui timol em seu óleo essencial, que se encontra armazenado em tricomas glandulares. O timol é um componente majoritário de importância econômica na indústria química e farmacêutica. O objetivo do trabalho foi estudar o efeito das temperaturas de secagem de folhas de L. origanoides na integridade dos tricomas, teor e composição química do óleo essencial. O delineamento experimental foi inteiramente casualizado, com quatro tratamentos definidos por temperaturas de secagem em estufa (40ºC, 50ºC, 60ºC e 70ºC) e quatro repetições. Foram realizadas extrações por hidrodestilação e determinação dos teores dos óleos essenciais (%), análise de composição química do óleo por cromatografia gasosa acoplada à espectrometria de massas e microscopia eletrônica de varredura para determinação da integridade dos tricomas. A secagem das folhas de L. origanoides em estufa de ventilação forçada a 40ºC minimizou a perda do teor de óleo essencial extraído (17,5g kg-1) e porcentagem relativa de timol. Folhas secas em temperatura de 60 e 70ºC exibiram maior porcentagem de tricomas rompidos e redução do teor de óleo essencial para 13,7g kg-1 e 11,8g kg-1, respectivamente.

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Histochemical analysis is essential for the study of plant secretory structures whose classification is based, at least partially, on the composition of their secretion. As each gland may produce one or more types of substances, a correct analysis of its secretion should be done using various histochemical tests to detect metabolites of different chemical classes. Here I describe some of the most used methods to detect carbohydrates, proteins, lipids, phenolic compounds, and alkaloids in the secretory structures.

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ABSTRACT Piper callosum Ruiz & Pav., Piperaceae, popularly known as “elixir-paregórico” and “matricá” in Brazil, is used in folk medicine to treat gonorrhea, general pain, and digestive disorders, and has repellent, astringent, diuretic, depurative, and haemostatic properties. Despite the fact that this plant is sold as a traditional phytotherapeutic product, we did not find reports on its quality control. We, therefore, performed macroscopic, microscopic, histochemical, and physicochemical analyses using standard methods to establish botanical authentication and purity degree parameters for leaves and stem of this species in two forms: medicinal plant and herbal drug. We observed the size, shape, color, texture, fracture surface and transection characteristics, leaf venation patterns, and calluses are valuable diagnostic characters to identify the herbal drugs when they are not ground or powdered. Since medicinal plants and herbal drugs did not differ anatomically, the following key anatomical characters for P. callosum can be used for diagnostic purposes of both types raw plant materials: epicuticular wax and cuticular flanges patterns; collenchyma features; fibers in the midrib; arrangement pattern of the vascular bundles of the midrib and petiole; shape of the midrib, leaf margin, petiole, and stem; occurrence of raphides; and morphology of the starch grains. Acid lipids, essential oils, oleoresins, steroids, tannins and flavonoids were histochemically identified. Total ash (leaves: 11.25%; stem: 5.25%), sulphated ash (leaves: 68.02%; stem: 12.50%), acid-insoluble ash (leaves: 2.82%; stem: 0.27%), moisture (leaves: 8.60%; stem: 6.10%), loss on drying (leaves: 11.08%; stem: 8.58%), and pH (leaves: 5.57, stem: 5.28) values were determined. The order of analyzed metal levels in leaf and stem herbal drugs was Al > V > Cu > Mn > Cr > Ni. Similar levels of Cd and Co and low levels of Hg were found. The results obtained can be used as quality control parameters for medicinal plants and herbal drugs of P. callosum.