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Licopoli 'organs' (Lc o) at epidermis level of the lamina of Statice lychnidifolia (st – stomata) [MAURY, 1886]  

Licopoli 'organs' (Lc o) at epidermis level of the lamina of Statice lychnidifolia (st – stomata) [MAURY, 1886]  

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Salt (chalk) glands of Plumbaginaceae represent interesting structures involved in the excretion of calcium carbonate outside plants' organs, especially on leaves surfaces. These chalk-glands, nominated by some authors as 'Licopoli' or 'Mettenius' organs are also very important from taxonomical point of view. Their structure has been a matter of de...

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... The function and structure of salt glands of the species of Plumbaginaceae have been reviewed (Grigore and Toma 2016;Caperta et al. 2020). The salt glands of this family consist of eight to 40 cells with the most frequent number of 16. ...
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In this study, we examined leaf anatomy of 24 species belonging to Acantholimon from flora of Iran. This paper, therefore, describes the foliar anatomical characters in details and discusses the extent to which leaf anatomical features might be developed in response to arid, semi-arid and mountainous condition of the Irano-Turanian region. The results revealed that the leaf anatomy of Acantholimon is highly consistent with the species habitat. The leaf anatomical characters such as the presence of salt glands and trichomes, isobilateral leaves, and sclerophylly are the most predominant responses to the environmental conditions. This paper is the first report of the existence of sclereids in Acantholimon species. The survey of the secreted elements from salt glands on the leaves epidermis of Acantholimon species were performed using SEM energy-dispersive X-ray (EDS) analysis for the first time. Based on these results, the crystals were determined to be truncated conical in shape, composed largely of Ca, O and C elements. Taxonomic implication of the anatomical characters in Acantholimon has been also discussed.
... He was the first who tried to analyse minerals secreted by these glands of different species of Satice and Plumbago. But his work remained unknown to many future botanists for many years (Grigore & Toma, 2016). ...
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Stem and leaves of two species of PlumbagoLinn.viz. P. zeylanica Linn.andP. auriculata Lam. were investigated for the structure and chemical composition of chalk glands. Light Microscopy (LM) and Scanning Electron Microscopy (SEM) revealed the presence of chalk glands on both lower as well as upper surface of leaf and stem of both species. Chalk glands are abundant on lower surface and sparse on upper surface of leaf. Chalk glands are approximately hemispherical glands with oval or almost circular outline. It is composed of 8 cells arranged in two circles – central circle of 4 secretory cells and outer circle of 4 adjoining cells. Each secretory cell has depression which corresponds to pore. Each gland is surrounded by 4 subsidiary cells. No significant difference in the structure of chalk glands in both species was noticed. Chalk glands occupy three different positions with regard to epidermal cells –at the same level of the epidermis, slightly sunken in the epidermis and slightly raised above the epidermis. Common elements found in EDAX analysis of all chalk glands are carbon, oxygen, magnesium, sulphur, potassium and calcium. Differences in the presence of elements silicon, chlorine, aluminium, sodium, phosphorus were observed. The presence of significant amount of calcium in chalk glands and their dried deposits and absence of sodium and chlorine from dried deposits and even in some chalk glands appealed to use the term ‘Chalk gland’ instead of ‘Salt gland’ in Plumbago.
... The morphology and anatomy of vegetative organs in Limonium species are generally well known and have been recently reviewed [23,72]. Studies on Limonium's biology have been mostly linked to the presence of salt glands ('Mettenius' or 'Licopoli' glands/ organs-see [73]), located on the stem and especially on the leaf surface. In fact, there are also some anatomical features found in Limonium that attest the xeromorphic nature of these halophytic species. ...
... This feature classifies halophyte species from Plumbaginaceae in the group of recretohalophytes (that secrete salt via salt bladders and salt glands-the case of Limonium species). Salt glands are particularly well-known and of great scientific interest from the historical (see [73] and references therein), anatomical [71,73,78,85], physiological [86] and evolutionary [87,88] points of view. Within the recretohalophytes from Plumbaginaceae, recent data report 48 species of Limonium to have salt glands [87]. ...
... This feature classifies halophyte species from Plumbaginaceae in the group of recretohalophytes (that secrete salt via salt bladders and salt glands-the case of Limonium species). Salt glands are particularly well-known and of great scientific interest from the historical (see [73] and references therein), anatomical [71,73,78,85], physiological [86] and evolutionary [87,88] points of view. Within the recretohalophytes from Plumbaginaceae, recent data report 48 species of Limonium to have salt glands [87]. ...
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Limonium is one of the most interesting and biodiverse genera of halophytes, with many species adapted to saline environments. Limonium species have a promising potential as cultivated minor crops as many have ornamental value, or are already used as medicinal plants. Other species are marketed as gourmet food or can be used for decontamination of polluted soils. Design and implementation of specific breeding programmes are needed to fully realise this potential, based on the vast genetic variation and high stress tolerance of wild species within the genus. Most Limonium species are halophytes, but many are also resistant to drought, especially those from the Mediterra-nean and other arid regions. Such species constitute attractive models for basic research on the mechanisms of stress tolerance, both constitutive and induced. As typical recretohalopyhtes, with excretive salt glands, Limonium species possess remarkable morpho-anatomical traits. Salt tolerance in this genus relies also on ion accumulation in the leaves, the concomitant use of diverse osmolytes for osmotic adjustment, and the activation of efficient antioxidant systems.
... However, in natural populations of G. italicum Tammaro, Pignatti & G.Frizzi, poor seed germination has been associated with the prevalence of vegetative over generative reproduction (Petriccione and Marrone 2013). Goniolimon taxa have more or less developed salt glands on aboveground organs (leaves and stems) that are common in plants that grow in arid and halophilous environments (Faraday and Thomson 1986;Grigore and Toma 2016). Although reports on chromosome counts in certain Goniolimon taxa are sometimes ambiguous, they reveal that the chromosome number in different representatives of this genus is 2n = 32 (Ančev 1982;Tammaro et al. 1982;Malakhova 1990), 2n = 34 (Runemark 1974;Strid, 2015) or 2n = 36 (Pignatti 1972). ...
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Goniolimon species are mainly components of the Eurasian steppe or steppe-like rocky vegetation, with some taxa occurring also in south-eastern Europe and northern Africa. We analysed the variability of: (i) two maternally inherited plastid loci (rpl32-trnL and 3′rps16-5′trnK) in 110 individuals of six currently accepted species from the Balkans and one species from the Apennines, to provide new insights into their origin and evolutionary history; and (ii) quantitative morphological characters (14 independent characters and one ratio character) in 641 individuals of three species of which two are morphologically and ecologically similar (G. italicum and G. tataricum) and the third, G. dalmaticum, was frequently misidentified as G. tataricum in the past, to provide new taxonomic treatment for proposed G. tataricum subspecies. We delineated several quantitative and five qualitative characters studied in a more limited sample as diagnostic for the identification of four subspecies (three newly described and one in a new rank) of G. tataricum. The history of westward peripheral populations of this species in the Balkans and the Apennines was rather complex and driven by local geo-historic events. These events facilitated multiple waves of east–west expansion of lineages originating from sources outside of the Balkan Peninsula which periodically diversified and occupied localised areas in the Balkans during the Pleistocene. An initial spread of an ancient G. tataricum lineage throughout south-eastern Europe probably occurred during the Messinian Salinity Crisis. Inter- and intraspecific hybridisation/introgression, as well as retention of ancestral polymorphisms, was common in G. tataricum and related taxa over time.
... Salt glands of P. auriculata are found on the adaxial and abaxial epidermal surfaces of the leaves and are known to excrete salt that form crystals on leaf surfaces (Ceccoli et al., 2015). These glands are common in the family Plumbaginaceae (Grigore and Toma 2016). In the 1800's these glands were initially referred to as chalk glands due to the secretion of insoluble carbonate salts (Metcalfe and Chalk 1972;Grigore and Toma 2016). ...
... These glands are common in the family Plumbaginaceae (Grigore and Toma 2016). In the 1800's these glands were initially referred to as chalk glands due to the secretion of insoluble carbonate salts (Metcalfe and Chalk 1972;Grigore and Toma 2016). Later it was named "salt glands" after it was discovered that these structures secrete NaCl (sodium chloride) (Faraday and Thomson, 1986a;Grigore and Toma 2016). ...
... In the 1800's these glands were initially referred to as chalk glands due to the secretion of insoluble carbonate salts (Metcalfe and Chalk 1972;Grigore and Toma 2016). Later it was named "salt glands" after it was discovered that these structures secrete NaCl (sodium chloride) (Faraday and Thomson, 1986a;Grigore and Toma 2016). The terms "salt glands and "chalk glands" tend to be interchangeable and to date no general distinctions have been drawn between the two terms (Dassanayake and Larkin 2017). ...
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This study interprets the structure and provides insights on the phyto-compounds of the leaves and calyces of Plumbago auriculata and evaluates its potential as a medicinal plant based on the presence of these compounds. Structure and possession of phyto-compounds were determined using light and electron microscopy and various histochemical tests. Glandular capitate and non-glandular uniserate trichomes were present on the surface of the calyx. Glandular trichomes bear a close resemblance to the digestive glands of the genus Drosera. Salt glands present on the leaf surface were abundant on the abaxial surface and revealed the presence of “transfer cells” reported for the first time in the genus. Abundant mitochondria, mini vacuoles, ribosomes, dictyosomes and rough endoplasmic reticulum cisternae were actively involved in the secretory process. Histochemical staining also revealed the presence of alkaloids and phenolic compounds that are of medicinal importance used to treat multiple ailments and also function as chemical deterrents in plants. Overall, this study contributes to the existing body of knowledge on the secretory structures of P. auriculata and provides new insights on the ultrastructural and histochemical aspects of the species.
... Density of salt glands observed in the present study was found to be much lower than the related taxa of plumbaginaceae described in Plumbago capensis, P.europaea and Limonium latifolium and Statice gmelin by Grigore and Toma. 18 In Limonium species of Plumbaginaceae it was reported to contained 20 to 50 salt glands per square millimeter of leaf. 19 ...
... Glandular secreting glands are common on the exterior surface of the calyx of flowers of Plumbaginaceae family. 7,18 In Plumbago zeylanica, glandular secreting glands was found distributed in between the nerves (Figure 8 and 8a) throughout the length of the calyx tube. Calyx indumentum was glabrous in nerves ( Figure 8). ...
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Background: Salt glands are highly specialized epidermal structures developed in recretoha-lophytic plants, intended for salt storage (reservoir) and as a means to alleviate the salt stress of the saline habitat by exo-recreto releases excess salts from the mesophyll tissues to the surrounding environment. Plumbago zeylanica Linn belongs to the family Plumbaginaceae is an important medicinal plant and espoused as a source for the drug Chitraka. Aim: The present study concerns the delineation of micromorphological characteristics of the paradermal and cross sections of the leaf epidermis, lamina, salt glands, petiole and mucilage secreting glands of the calyx of the flowers of Plumbago zeylanica Linn. Materials and Methods: Delimiting the morpho-histological profile of the leaves and to develop the chromatogram of the methanol extract of the leaves and quantification of the marker compound plumbagin in the leaves using High Performance Thin- Layer Chromatographic (HPTLC) method. Results: Distribution of uniseriate, cruciate type clothing trichomes, characteristic contour of midrib, presence of amphistomatic epidermis with anisocytic stomata, crystals in idioblast of ground tissues of petiole and mesophyll, large palisade ratio, small stomatal index, multicellular salt gland structure with 8 cells, cuticular secretory cells, crescent shaped petiole, densely distributed mucilage secreting glands on the calyx of flowers were features characteristic of the taxon. The HPTLC finger printing profile of the leaves revealed six phyto-constituents. Densitometric scanning of plumbagin had shown the absorption spectra λmax at 270 nm. Spectral matching by overlaying the absorption spectra of standard marker compound with the phyto-constituents present in the methanol extract of sample could fail to obtain any spectral matching at λmax 270 nm, suggestive that dried leaves of P. zeylanica did not contain any appreciable amount of Plumbagin. Calibration data obtained by polynomial regression had revealed that dried leaves of P. zeylanica Linn contained below 105 ng/μl of plumbagin in methanol extract of leaves.
... Literature reports that salt glands occur sunken in the epidermis of leaves and are associated with ion secretion whereas trichoms protrude from the surface of leaves or stems and are often responsible for the secretion of exudates (Cutter, 1978, Fahn, 1988Naidoo and Naidoo, 1998). Although some researchers report salt glands to be specialized trichomes, based on available literature for the purpose of this review salt glands and trichomes are classified independently as secretory structures in Plumbaginaceae ( Metcalfe and Chalk, 1972;Thomson, 1986b, Grigore andToma, 2016). a b c d ...
... Salt glands of the family Plumbaginaceae are multicellular and sunken in the leaf epidermal cells, consisting of basal and secretory cells with the cells varying in number from six up to forty (Thomson, 1975, Faraday andThomson, 1986b). Glands were first described in the 1800's and were referred to as chalk glands due to the presence of insoluble carbonate salts found above the gland on the surfaces of stems and leaves (Grigore and Toma, 2016) . However, when it was later discovered that the glands of some species secrete sodium chloride (NACl) these glands were then referred to as salt glands ( Faraday and Thomson, 1986a;Grigore and Toma, 2016). ...
... Glands were first described in the 1800's and were referred to as chalk glands due to the presence of insoluble carbonate salts found above the gland on the surfaces of stems and leaves (Grigore and Toma, 2016) . However, when it was later discovered that the glands of some species secrete sodium chloride (NACl) these glands were then referred to as salt glands ( Faraday and Thomson, 1986a;Grigore and Toma, 2016). The salt glands (Mettenius glands or Licopoli glands), as described by Grigore and Toma (2016), based on historical facts by Metcalfe and Chalk (1972) (Figure 3c) (Faraday and Thomson, 1986a). ...
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Background: The genus Plumbago distributed in warm tropical regions throughout the world is the largest genus in Plumbaginaceae. Medicinal plants are characteristic to the genus Plumbago and are cultivated and utilized worldwide. Plumbago auriculata Lam. is common in South Africa and is often cultivated for its ornamental and medicinal uses throughout the world. Materials and Methods: A comprehensive review of the genus Plumbago with focus on Plumbago auriculata was carried out and information was gathered using scientific publications, conference proceedings, the internet and books. Articles based on the morphology, pharmacological and medicinal uses of Plumbago auriculata was analysed thoroughly. Results: Plumbago auriculata plant parts posses a wide range of phytochemicals with plumbagin being the marker compound showing various pharmacological activities. Different plant parts are claimed to be used for the treatment of human and animal ailments, however they do exhibit toxic properties and need to be administered with caution. Salt secreting glands and trichomes are characteristic of Plumbaginaceae. Conclusion: This study reveals new insights on the genus Plumbago and the potential use of species in the genus as medicinal plants. Plumbago auriculata possess the bioactive compound plumbagin and secondary metabolites, thus, it is of high medicinal importance. P. auriculata is a poorly nor favourite studied species in the genus Plumbago and further research needs to be carried out to explore specific details of the species.
... (Naugolnykh & Kerp 1996, Plate V, Figs 9, 10). Similar concentrically organized glands are known for the recent angiosperms of the family Plumbaginaceae (Grigore & Toma 2016). Normally, secretory glands are much smaller, but somewhat larger multicellular glands and extrafloral nectaries are also known (Diaz-Castelazo et al. 2005), and some of them have similar shape (Diaz-Castelazo et al. 2005;Figs 2E, 3B, 4B;Machado et al. 2008;Sawidis 2012, Fig. 5) and can be arranged in a similar dense pattern (Diaz-Castelazo et al. 2005, Fig. 2F) although they are still quite small. ...
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A new concept and a newly emended diagnosis of the advanced peltasperm Kuvakospermum pedatum Naug. et Sidorov, emend. nov. from the Kazanian (Middle Permian) deposits of Russia Serge V. Naugolnykh Summary: The paper is devoted to the monotypic genus Kuvakospermum Naug. et Sidorov, emend. nov., which is interpreted as a highly advanced representative of peltasperms (order Peltaspermales). The collection studied includes six specimens of Kuvakospermum pedatum Naugolnykh et Sidorov, emend. nov. (four of them are practically completely preserved megasporophylls and two specimens with the seeds preserved in natural connection to reproductive organ, but with the partly or completely detached megasporophyll shield). The megasporophyll is stalked, peltate, with round to ovoid megasporophyll shield with the lobed margin bearing long linear extenuations. Adaxial surface of the megasporophyll shield bears well-developed protective belt. The seeds were attached to adaxial surface of the megasporophyll in the space between the protective belt and the stalk, disposed concentrically around the stalk. Adaxial surface of the megasporophyll bears small secretory glands forming regular parastichies. Abaxial surface of the megasporophyll shield bears numerous small round resin bodies. Some possible fossil relatives and recent morphofunctional analogues of Kuvakospermum pedatum are discussed.