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Main components (>5%) of the essential oils of Helichrysum stoechas previously studied 

Main components (>5%) of the essential oils of Helichrysum stoechas previously studied 

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The aerial organs of Helichrysum stoechas bear a dense woolly indumentum with two types of trichomes: nonglandular trichomes, extremely long, simple, uniseriate and multicellular; and biseriate, glandular trichomes with 10-14 cells. Glandular trichomes are abundant on the leaf abaxial surface, particularly on the intervein areas, on the inflorescen...

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... (Moench), respectively. In the oil analyzed by Chinou et al. (1997a,b), β-caryo- phyllene (16%) and β-elemene (13%) were the domi- nant components, and in the oil analyzed by Tsoukatou et al. (1999), α-pinene (28%) and epi-α-bisabolol (22%) were the main ones, whereas α-pinene (63%) was the major component of H. stoechas oil studied by Vernin and Poite (1998) (see Table 2). The results of the latter study, together with those of Proença da Cunha and Relative to C 8 -C 17 n-alkanes on the DB-1 column. ...

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In the present work three subspecies of Helichrysum plicatum DC. (Helichrysum plicatum DC. subsp. plicatum, Helichrysum plicatum DC. subsp. polyphillum (Ledeb) P.H.Davis & Kupicha and Helichrysum plicatum DC. subsp. isauricum Parolly) were investigated for the essential oil chemical compositions. The volatiles were obtained by conventional hydrodis...

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... Similarly to other Asteraceae species (Werker and Fahn, 1981;Werker et al., 1994;Ascensão et al., 2001;Sulborska, 2013), the highest glandular trichome density is typical of the abaxial side of the I. britannica leaf (Fig. 3A). According to Maffei et al. (1989) the rate of trichome differentiation varies on the adaxial and abaxial sides of the leaf. ...
... The morphology (number of cells, their size, and shape) of the biseriate cylindrical trichomes of I. britannica on the leaves is fairly uniform and homogeneous. Trichomes of different morphology and functions, from non-glandular to glandular, have been observed in the species of the Asteraceae family (Werker and Fahn, 1981;Werker et al., 1994;Ascensão et al., 2001;Sulborska, 2013;Viera Barreto et al., 2016;Bombo et al., 2016). Generally, several types of glandular trichomes were perceived in the species of family Asteraceae: uniseriate with a multicellular head and a unicellular base, uniseriate with a unicellular head and a multiseriate base, biseriate vesicular, biseriate pedunculate, sessile and stalk trichomes (Werker and Fahn, 1981;Krak and Mráz, 2008;Sulborska, 2013;Fernandes et al., 2016). ...
... Generally, several types of glandular trichomes were perceived in the species of family Asteraceae: uniseriate with a multicellular head and a unicellular base, uniseriate with a unicellular head and a multiseriate base, biseriate vesicular, biseriate pedunculate, sessile and stalk trichomes (Werker and Fahn, 1981;Krak and Mráz, 2008;Sulborska, 2013;Fernandes et al., 2016). Biseriate glandular trichomes are typical and present in several species of the Astereaceae family (Werker and Fahn, 1981;Duke and Paul, 1993;Ascensão et al., 2001;Ciccarelli et al., 2007;Dosa, 2009: Fernandes et al., 2016Liesenfeld et al., 2019;Muravnik et al., 2019). ...
Article
Glandular trichomes are secretory organs that vary greatly in size, shape, location, type, and composition of secondary metabolites that are synthesized in them. They represent a protective chemical barrier and metabolic factories for the synthesis of bioactive compounds. We evaluated the effect of sucrose on the growth and biomass production, as well as on the distribution, density, and chemical composition of leaf glandular trichomes developed in vitro cultured Inula britannica, an important medical and aromatic species. Nodal segments (5 mm) were cultivated on the basal Murashige and Skoog medium with different concentrations (0 M, 0.01 M, 0.06 M, 0.1 M and 0.3 M) of sucrose. The lack of sugar does not retard growth, while a high sugar concentration inhibits biomass accumulation in vitro. The density and number of leaf glandular trichomes are changed with the medium composition variation. Morphoanatomical and histochemical analyses of the trichomes of the cultured I. britannica applying SEM, conventional light, fluorescent and Raman microscopy revealed the presence of nonglandular and biseriate glandular trichomes on the leaf surface. The histochemical analysis proved that glandular trichomes synthesized a complex mixture of biomolecules. The Raman microscopy analysis of glandular trichomes confirmed the presence of terpenes, most probably the bicyclic monoterpenes and sesquiterpenes. The multivariate analysis, based on PCA, was applied to distinguish among the glandular trichomes of three sugar concentrations: sugar-free medium (0 M), optimal (0.1 M) and the highest sucrose concentration (0.3 M). The main differences between the chemical profile of the leaf glandular trichomes of the plants grown on the 0 M and the chemical profile of the trichomes developed on the 0.1 and 0.3 M medium come mainly from phenolic compounds and to a smaller degree from cyclic terpenes, while the chemical profile of the glandular trichomes grown on the 0.1 M and 0.3 M differs considerably from the trichomes grown on the sugar-free medium in the polysaccharide cell wall ingredients. Most differences between the chemical profile of the glandular trichomes developed on 0 M and 0.3 M and the chemical profile of the trichomes grown on the 0.1 M of sucrose, come from phenolic compounds.
... Helichrysum stoechas (L.) Moench is a Mediterranean plant belong to the Asteraceae family, which contains almost 600 species, and the Helichrysum genus is one of them (Benelli et al., 2018). This species is an evergreen aromatic grey-cotton subshrub growing on sandy or shallow soils (Ascensão et al., 2001). Ethnobotanical surveys revealed that H. stoechas have been used to treat flu, common cold, wound healing, digestive disorders and many other medical issues (Barroso et al., 2014;Zengin et al., 2020). ...
Article
The present study was aimed to investigate the phenolic composition, antioxidant and anti-inflammatory activities of ethyl acetate (EAE) and n-butanol (BuE) extracts obtained from the aerial parts of Helichrysum stoechas (L.) Moench. Both extracts are rich in polyphenols and flavonoids with a wealth more marked for EAE. The LC-ESI-MS analysis revealed the presence of many important phenolics such as isoquercitrin, rutin, ferulic acid and chlorogenic acid. The extracts exhibited anti-inflammatory effect by inhibiting the croton oil-induced ear œdema and the neutrophil migration in carrageenan-induced air pouch model. These extracts stabilized erythrocytes membrane and inhibited protein denaturation. In addition, both extracts exerted antioxidant effect by scavenging superoxide anion radicals, hydroxyl radicals and hydrogen peroxide. Moreover, they exerted a significant protective effect against AAPH-induced erythrocyte hemolysis. Results obtained in the present study indicate that H. stoechas can be a potential source of natural antioxidant and anti-inflammatory agents.
... The occurrence of papillae only in the apical zone of corolla lobes and the absence of trichomes on the corolla were observed by Haratym and Weryszko-Chmielewska (2012). However, biseriate glandular trichomes were detected on corolla lobe tips in Helichrysum (Ascensão et al. 2001), whereas biseriate trichomes were observed on different corolla parts in Chamomilla and Inula (Sulborska 2011(Sulborska , 2013. ...
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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.
... One of the largest angiosperm family, Asteraceae, includes hundreds of species of economic interest, e.g., medicinal plants which are used therapeutically, and consequently, their anatomy and morphology have been studied, including the structures which produce and storage specialized metabolites (Werker and Fahn 1981;Ascensão and Pais 1987;Figueiredo and Pais 1994;Ascensão et al. 2001;Andreucci et al. 2008;Simpson 2009;Camilotti et al. 2014;Gavrilović et al. 2016). These secretory structures in Asteraceae could be found in the form of glandular trichomes, cavities, canals, laticifers, hydathodes, idioblasts and glandular appendices (Metcalfe and Chalk 1950;Carlquist 1958;Lestern and Curtis 1985;Castro et al. 1997;Gavrilović et al. 2018a). ...
Article
Comparative anatomical analysis of vegetative organs has been conducted on Artemisia campestris L., A. absinthium L., A. arborescens L., A. judaica L. and A. herba-alba Asso, using light microscopy, in order to examine the most important anatomical features and to find new valid taxonomic characters. Results have shown that general root, stem and leaf anatomical features and nonglandular and glandular trichomes are shared by all species. However, some characters (parenchyma sheath, which surrounded vascular bundle and extended to both epidermises, subepidermal collenchyma and the absence of secretory canals in the leaves) link together A. absinthium and A. arborescens from the same section. Some characters, as periderm and lignified pith parenchyma cells (A. campestris and A. arborescens), nonendodermal secretory canals in root cortex (A. absinthium and A. judaica) and secretory canals in the leaf phloem (A. judaica and A. herba-alba), connect species belonging to different sections. Moreover, some characters could be considered as species-specific, nonendodermal secretory canals in the root secondary phloem, triangular leaf shape on the cross section and secretory canals in the leaf parenchyma for A. campestris, secretory canals in the stem pith for A. absinthium, crystals in the pith parenchyma cells for A. arborescens and the absence of root secretory canals for A. herba-alba. Given results revealed qualitative characters, on the basis of which the studied species are anatomically distinguishable between each other, provide valuable features for better species identification and contribute to the anatomy of the genus Artemisia.
... Achenes were mounted with one drop of Tween 20 and one drop of Hoyer's Solution was added. In order to characterize all species included in the phylogenetic analysis, we complemented the data with information from the literature, especially for the species outside the Lucilia group, or where achenes were not available for study (Cabrera, 1932;Dillon & Sagástegui Alva, 1986, 1991aAnderberg & Freire, 1990;Anderberg, 1991;Freire, 1993Freire, , 1995Ascensão & al., 2001;GalbanyCasals & al., 2004a;Morefield, 2006;Abid & Qaiser, 2008b;Loeuille & al., 2011;AndrésSánchez & al., 2014AndrésSánchez & al., , 2015Urtubey & al., 2016). Appendix 2 contains a list of sources from which information on achenial trichome morphology was obtained. ...
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The Gnaphalieae (Asteraceae) are a cosmopolitan tribe with around 185 genera and 2000 species. The New World is one of the centers of diversity of the tribe with 24 genera and over 100 species, most of which form a clade called the Lucilia-group with 21 genera. However, the generic classification of the Lucilia-group has been controversial with no agreement on delimitation or circumscription of genera. Especially controversial has been the taxonomic value of achenial trichomes and molecular studies have shown equivocal results so far. The major aims of this paper are to provide a nearly complete phylogeny of the Lucilia-group at generic level and to discuss the evolutionary trends and taxonomic significance of achenial trichome morphology. We conducted a phylogenetic analysis of the New World Gnaphalieae with nrDNA (ETS, ITS) sequence data from a sampling of 18 genera of the Lucilia-group and utilized these results to examine morphological evolution of achenial trichome types and presence of apical myxogenic cells. Seven well-supported subclades can be recognized within the Lucilia-group (L1–L7). These results support Brazilian and Andean Berroa, Facelis, Lucilia, and Micropsis forming a clade (L1), the inclusion of Chilean Lucilia under Belloa (L2), the monophyly of Stuckertiella +Gamochaeta +Gamochaetopsis (L3), Chevreulia +Cuatrecasasiella (L4) and Antennaria (L5) excluding Antennaria linearifolia, which is resolved in a monophyletic group together with Jalcophila, Loricaria and Mniodes (L6), and the recognition of Gnaphaliothamnus (L7) removed from Brazilian taxa of Chionolaena (L2). Ancestral character state reconstruction of achenial trichome morphology suggests that clades are homogeneous in terms of trichome type, but with exceptions that make it highly homoplastic. Conversely, our results suggest that the presence of myxogenic apical cells is less homoplastic and that closely related species tend to resemble each other more than expected under random variation.
... Asteraceae include many species of economic interest, many of these are used therapeutically, and consequently their morphology and anatomy have been studied, including the structures responsible for the production, storage and secretion of secondary metabolic compounds (Werker & Fahn, 1981;Ascensão & Pais, 1987;Figueiredo & Pais, 1994;Ascensão et al., 2001;Andreucci et al., 2008;Camilotti et al., 2014). These secretory structures in Asteraceae take the form of glandular trichomes, canals, cavities, laticifers, idioblasts, hydathodes and glandular appendices (Metcalfe & Chalk, 1950;Carlquist, 1958;Lestern & Curtis, 1985;Castro, Leitão Filho & Monteiro, 1997). ...
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Vernonieae are well represented in South America by subtribe Lychnophorinae, with 104 Brazilian species, some of them rare and endangered. Lychnophorinae are well known for producing metabolites of high pharmaceutical and chemotaxonomic value. Despite the importance of secondary metabolites in Lychnophorinae, there is still little evidence regarding the secretory structures responsible for producing these compounds. Therefore, in this study we investigated 15 species representative of the main lineages of Lychnophorinae, with the following objectives: to identify where the chemotaxonomic markers of Lychnophorinae are secreted and stored and in which developmental stage of the organ these metabolites are found. Samples of stems and leaves were processed according to the usual techniques in plant anatomy. It was found that the main sites of secondary metabolite biosynthesis are glandular trichomes, epidermal cells and parenchyma tissues. Metabolites from glandular trichomes, especially sesquiterpene lactones (STL), are prevalent in the early developmental stages of organs. The metabolite compounds stored in parenchyma tissues are mainly terpenoids, flavonoids and other phenolic compounds; young and expanded leaves are equally rich in metabolites. Thus, the information obtained in this study is essential for conducting chemotaxonomic studies in this group, helping to promote selective collection and conservation of species.
... Among the aromatic plants of the Mediterranean area, Helichrysum species (Asteraceae, Gnaphalieae) are well-known for their secondary metabolite content, specifically flavonoids, sesquiterpene lactones and terpenes occurring in essential oils (Angioini et al., 2003;Ascensão et al., 2001;Bianchini et al., 2001Bianchini et al., , 2003Kladar et al., 2015;Leonardi et al., 2013;Mastelic et al., 2005;Melito et al., 2013;Morone-Fortunato et al., 2010;Ornano et al., 2015;Paolini et al., 2006;Perrini et al., 2009;Ruberto et al., 2002;Tsoukatou et al., 1999;Vernin and Poite, 1998;Viegas et al., 2014;Zeljkovi c et al., 2015). Helichrysum extracts are used in cosmetic industry and have potential pharmacological applications for their antioxidant, antimicrobial, and anti-inflammatory activities (Bremner, 2009;Lall, 2006;Mancini et al., 2011;Viegas et al., 2014). ...
... As a whole, HS-SPME analysis across Helichrysum populations revealed the production of nearly 400 different compounds, belonging to several chemical classes, whereas the total volatile compounds detected in hydrodistillated essential oils from several Mediterranean species ranged between 80 and 110: terpenes prevailed, hydrocarbons and alcohols were scarcely represented, esters were absent (Angioini et al., 2003;Ascensão et al., 2001;Bianchini et al., 2001Bianchini et al., , 2003Kladar et al., 2015;Leonardi et al., 2013;Mastelic et al., 2005;Melito et al., 2013;Morone-Fortunato et al., 2010;Ornano et al., 2015;Paolini et al., 2006;Perrini et al., 2009;Ruberto et al., 2002;Viegas et al., 2014;Tsoukatou et al., 1999;Vernin and Poite, 1998;Zeljkovi c et al., 2015). In particular, HS-SPME analysis detected a very higher number of terpenes (about 50% more) than hydrodistillation did, however displaying a general consistency with the literature data. ...
... The terpenic fraction of H. stoechas VOCs showed a general qualitative consistency with the available data on leaf essential oils (Ascensão et al., 2001;Tsoukatou et al., 1999;Vernin and Poite, 1998), a-pinene being invariably the dominant component. ...
Article
The species of Helichrysum sect. Stoechadina (Asteraceae) are well-known for their secondary metabolite content and the characteristic aromatic bouquets. In the wild, populations exhibit a wide phenotypic plasticity which makes critical the circumscription of species and infraspecific ranks. Previous investigations on Helichrysum italicum complex focused on a possible phytochemical typification based on hydrodistilled essential oils. Aims of this paper are three-fold: (i) characterizing the volatile profiles of different populations, testing (ii) how these profiles vary across populations and (iii) how the phytochemical diversity may contribute in solving taxonomic problems.
... Furthermore, the elimination of secondary metabolites by glandular trichomes may provide chemical protection against herbivores and pathogens (Werker 2000, Glas et al. 2012). The biseriate vesicular glandular trichomes of T. cinerea are similar to those found on the aerial organs of other Asteraceae species and several aspects of these structures have been investigated (Ascensão and Pais 1987, Figueiredo and Pais 1994, Afolayan and Meyer 1995, Pagni 1995, Castro et al. 1997, Ascensão et al. 2001, Heinrich et al. 2002, Andreucci et al. 2008, Oliveira et al. 2013, Trindade et al. 2014. ...
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Trichogonia cinerea is endemic to Brazil and occurs in areas of cerrado and campo rupestre. In this study, we characterized the glandular and non-glandular trichomes on the aerial parts of this species, determined the principal events in the development of the former, and identified the main constituents of the volatile oil produced in its aerial organs. Fully expanded leaves, internodes, florets, involucral bracts, and stem apices were used for the characterization of trichomes. Leaves, internodes, florets, and involucral bracts were examined by light microscopy and scanning electron microscopy, whereas stem apices were examined only by light microscopy. Branches in the reproductive phase were used for the extraction and determination of the composition of the volatile oil. The species has three types of glandular trichomes, biseriate vesicular, biseriate pedunculate, and multicellular uniseriate, which secrete volatile oils and phenolic compounds. The major components identified in the volatile oil were 3,5-muuroladiene (39.56%) and butylated hydroxytoluene (13.07%).
... According to Werker and Fahn (1981), glandular trichomes limit the transpiration rate and reduce leaf temperature. Withering and falling leaves with intact glandular hairs may also provide a phytotoxic environment for germinating seeds and growing seedlings while non-glandular trichomes often act as a physiological barrier against herbivores and contribute to plant adaptation to environmental conditions, particularly in dry environments ( Afolayan and Meyer, 1995;Ascensao et al., 2001;Aneta, 2013). These attributes may explain the wide distribution of L. javanica in various climatic environments in Southern Africa. ...
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Background: Lippia javanica (Burm.F.) Spreng is an aromatic indigenous South African plant with culinary and medicinal values. This study investigated the foliar morphology and elemental composition of the plant because not much data concerning the anatomical and micro-morphological features can be found in literatureMaterials and Methods: Fresh leaves of the plant were investigated using light and scanning electron microscopy (SEM). The elemental composition of the leaf was determined by energy dispersive X-ray spectroscopy.Results: The leaves of L. javanica were amphitrichomic while the stomata distribution was amphistomatic. These stomata were more or less randomly distributed over the epidermis lying almost close to each other and were fewer in number in between the veins and over the finer veins. The major constituents of crystals found in the plant were Ca, Na, S, Al, P, Cl, K, Mg and Fe. The presence of long glandular trichomes on the leaf surfaces of the plant could indicate secretory sites where secondary metabolites are produced.Conclusion: Secretions from the glandular trichomes and the presence of these elements, which are vital in maintaining good health, are probably responsible for the culinary and medicinal properties of L. javanica.
... Results showed that both species present two types of secretory structures: glandular trichomes-capitate wit3 h a biseriate-pluricelular head, and secretory ducts, confirming the observations of Delbón et al. [36] [37]. Biseriate glandular trichomes are typically found in Asteraceae, in different tribes and genera, such as Ambrosia trifida [44], Helianthus annuus [45] [46], Sigesbeckia jorullensis [41], Grindelia pulchella/Heliantheae [29], Centrapalus pauciflorus and Chrysolaena spp./Vernonieae [28] [47], respectively, Helichrysum [48] and Inula/Inuleae [49], Artemisia annua [50] and Tanacetum cinerariifolium/Anthemideae [51]. ...
... Storage of secretions in the subcuticular space and cuticle rupture has been reported for other Asteraceae species [28] [29] [41] [48] [49] [58]. On the other hand, Fahn [55] reports that secretion by the apical head cells continue as long as the cells on the peduncle remains functional, which might be the case for FO. ...
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In this work, the localization, density, morphology and ultrastructure of secretory structures in aerial organs of Flourensia campestris (FC) and F. oolepis (FO) (Asteraceae) by means of a combination of light, fluorescence, transmission (TEM) and scanning electron microscopy (SEM) were examined. The possible role of secretory structures in the production and secretion of the phyto-toxic sesquiterpene (-)-hamanasic acid A ((-)HAA) in both species was also assessed. Capitate glandular trichomes were found in all reproductive organs of FC and FO, and were being reported for the first time. These glandular trichomes, typically associated to edges and veins, were of the same type as those already described for vegetative organs, and were abundant in involucral bracts and corolla of tubulose and ligulate flowers. Their density in reproductive organs of both species was similar (ca. 30/mm 2) and lower than that found in leaves (ca. 100/mm 2) and stems (ca. 160/mm 2 in FC, and up to 650/mm 2 in FO). Glandular trichomes in vegetative organs followed a species-* Corresponding authors. M. P. Silva et al. 926 specific pattern of distribution. TEM and SEM observations suggest that each species differs in the way in which secretory materials are released to the outside: through cracks or pores in FC, or through a loose cuticle in FO. Similar inspections of the secretory ducts revealed lipophilic vacuoles localized in subepithelial and epithelial cells, in which secretions accumulated before being transferred to the duct. The presence of wall ingrowths in subepithelial cells suggests that granulocrine secretion operates in these species. Secretory ducts varied in density and diameter among the organs in both species, with the combination being maximal in woody stems. (-)HAA was only detected in surface secreted resins of both species, and its concentration (2D-TLC, GC-FID) was intimately associated with the distribution and density of glandular trichomes in each organ (capitula, leaves, and stems with primary or secondary growth). In addition, no (-)HAA was detected internally in the resins collected from secretory ducts. The composition of these resins showed distinctive profiles for FC and FO, and only four from ca. 30 compounds detected (GC/MS) were shared by both species. In addition to the elucidation of ultrastructural traits, distribution and density of secreto-ry structures in aerial organs of FC and FO, present findings suggest a functional role for glandular trichomes in the secretion of the putative phytotoxic allelochemical (-)HAA.