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Swertia

Swertia is a genus of flowering plants in the family Gentianaceae, comprising 166 accepted species of annual and perennial herbs primarily distributed in temperate, subalpine, and mountainous regions of the Northern Hemisphere and Old World tropics.^[1] These plants are characterized by erect stems, opposite or rarely whorled leaves, and 4- to 5-merous flowers arranged in racemes, panicles, or solitary, featuring rotate corollas with short tubes, nectariferous lobes often adorned with fimbriae or scales, and oblong capsules containing winged or unwinged seeds.^[2] The genus exhibits significant diversity, with the highest number of species (around 100) occurring in Asia, followed by Africa (approximately 30 species), while Europe, North America, and Madagascar each host fewer species (3, 1, and 1, respectively).^[2] Habitats typically include open areas such as marshes, grasslands, and meadows in temperate zones.^[2] Several Swertia species hold notable economic and medicinal value, particularly in traditional systems like Ayurveda and Unani medicine, where they are used for their bitter secoiridoid compounds such as amarogentin and swertiamarin, which exhibit hepatoprotective, antidiabetic, antimicrobial, and antimalarial properties.^[3] For instance, Swertia chirayita, a key species from the temperate Himalayas, is employed as a tonic for liver disorders and has been included in historical pharmacopeias for its therapeutic potential.^[3] Conservation efforts are underway for some species due to overexploitation and habitat loss in regions like the Indian Himalayan Region, where over 32 species occur.^[4]

Taxonomy and phylogeny

Etymology and history

The genus name Swertia derives from Emanuel Sweert (also spelled Sweerts or Swert, 1552–1612), a Dutch botanist, florist, engraver, and author of the illustrated herbal Florilegium (1612), which featured detailed depictions of plants; Carl Linnaeus honored him by establishing the genus in this Linnaean tradition.^[2]^[5] Linnaeus first described Swertia in his Species Plantarum (1753), including five species primarily from temperate regions of Europe and Asia, marking the formal recognition of the genus within Gentianaceae.^[6] Early taxonomic accounts, such as those by Borkhausen (1796), often confused Swertia with the closely related genus Gentiana owing to shared traits like opposite leaves and tubular corollas, leading to initial misclassifications of several species.^[6] During the 19th century, botanist August Heinrich Rudolf Grisebach advanced the genus's classification by establishing the subtribe Swertiinae in 1837 and providing detailed species accounts in his 1838 work on Gentianaceae, which expanded the known diversity and clarified boundaries.^[6] This era saw refinements through segregation of allied genera like Lomatogonium and Pleurogyne from Swertia, alongside resolution of synonyms such as Exacum rosulatum Baker (now Swertia rosulata), contributing to the delimitation of approximately 137 species as recognized in 2015.^[6]^[7] In European cultural history, Swertia species, especially S. perennis known as felwort, appear in medieval and early modern herbals for their medicinal value as bitter tonics to alleviate digestive disorders and fevers, akin to uses of related gentians in traditional pharmacology.^[8]

Classification and synonyms

Swertia is classified within the family Gentianaceae, order Gentianales, in the tribe Gentianeae and subtribe Swertiinae.^[1]^[9] The genus shows close phylogenetic relationships to other members of subtribe Swertiinae, including Frasera and Gentianella, based on shared morphological and molecular traits such as corolla morphology and plastid gene sequences.^[10]^[11] Molecular phylogenetic analyses, particularly those employing nuclear internal transcribed spacer (ITS) regions and chloroplast matK gene sequences, have demonstrated that Swertia is polyphyletic, with its species nested in multiple clades alongside genera like Bartonia, Halenia, and Gentianopsis.^[12]^[10] These findings have prompted taxonomic revisions to establish monophyletic groups, including the segregation of certain lineages into genera such as Veratrilla, while retaining Swertia sensu stricto for a core group of Eurasian species.^[13]^[11] However, a 2025 phylotranscriptomic study proposes resolving the polyphyly by merging multiple genera (including Bartonia, Comastoma, Frasera, Gentianella, Halenia, Jaeschkea, Lomatogonium, and Veratrilla) into an expanded monophyletic Swertia, potentially encompassing approximately 500 species; this revision remains a proposal and has not yet been adopted in major databases like POWO.^[14] Historically, several genera have been treated as synonyms of Swertia due to overlapping morphological features, including rotate or tubular corollas, nectariferous scales, and calyx structure; these include Frasera Walter (established for North American species with whorled leaves), Lomatogonium A. Juss. (for annuals with solitary flowers), and Pleurogyne Eschsch. (for species with plicate corolla lobes).^[15]^[16]^[17] Such synonymy reflects early classifications based on vegetative and reproductive similarities, though modern phylogenomics supports their distinction in many cases.^[9] The genus encompasses approximately 166 accepted species worldwide as of November 2025 (Plants of the World Online), though recent proposals suggest potential expansion to over 500 via generic mergers.^[1]^[18] The type species is Swertia perennis L., a perennial herb native to Europe and northern Asia, designated under the genus's original description in 1753.^[19]

Description and biology

Morphology

Swertia species are primarily annual or perennial herbs, typically growing to heights of 10–100 cm, with erect, simple or branched stems that are terete, striate, or quadrangular in cross-section. The stems often bear a brownish-purple hue in some species and can reach up to 1.5 m in length, though most are shorter and herbaceous in nature.^[20]^[2]^[3] Leaves in the genus are arranged oppositely, rarely alternately or in whorls, and are entire-margined, sessile or amplexicaul, with shapes ranging from lanceolate to ovate and lengths up to 4 cm. They feature prominent 3–7 dorsal veins and basal leaves that may be petiolate and deciduous, while cauline leaves clasp the stem; the foliage imparts a characteristic bitter taste attributable to the presence of secoiridoid glycosides concentrated in the tissues.^[20]^[2]^[3] The root system consists of fibrous or woody primary roots with few secondary rootlets, or short rhizomes bearing fleshy adventitious roots, supporting the erect habit. Inflorescences are cymose, forming simple or paniculate thyrses, racemes, or solitary terminal flowers on often elongate pedicels. Flowers are 4–5-merous (rarely 6–7), with a rotate calyx and corolla that has a very short tube less than 3 mm long and lobes to the base; the corolla is typically blue-purple, though varying to greenish-yellow with purple tinges in species like S. chirayita, where individual flowers measure 1–3 cm in diameter, featuring 1–2 basal nectaries per lobe often fringed with fimbriae or scales.^[20]^[2]^[3] Fruits are oblong to ovoid capsules that dehisce septicidally via two valves, containing numerous small, subglobose to compressed seeds that are smooth, light brown or grey, and often winged or bearing ridge-like outgrowths for dispersal.^[20]^[2]

Reproduction and life cycle

Swertia species exhibit diverse reproductive strategies adapted to their temperate and montane habitats, with flowering typically occurring from late summer to autumn. For instance, in Swertia chirayita, flowering begins in August and continues through November in the third year of growth, while S. bimaculata displays protandrous dichogamy, with male-phase anthers dehiscing first to facilitate cross-pollination before the female phase.https://www.tandfonline.com/doi/full/10.1080/00087114.2013.780436^[21] This temporal separation reduces self-pollination, though some species like S. bimaculata show herkogamy in the female phase due to stamen retraction, further promoting outcrossing.^[21] Pollination in Swertia is predominantly entomophilous, relying on insects such as bees, syrphid flies, and calliphorid flies as primary vectors. Nectar-producing nectaries guide pollinators via distinct tracks on the corolla, enhancing visit efficiency and pollen transfer. Many species are self-compatible, as seen in S. chirayita, where autogamy yields 34.8% fruit set and geitonogamy 44.2%, though open pollination achieves up to 91.6% success, indicating a preference for xenogamy.^[21]^[22] Seed dispersal occurs primarily through anemochory, facilitated by winged seeds produced in ovate capsules containing up to 50 seeds per fruit, as in S. perennis. Germination in temperate species often requires cold stratification to break dormancy; many exhibit morphophysiological dormancy due to immature embryos. For example, S. thomsonii seeds achieve 84-86% germination after 20 days of wet stratification at 4°C, combined with chemical treatments like gibberellic acid.^[23]^[24] Life cycle variations reflect habitat demands, with pluri-annual (monocarpic perennial) species like S. chirayita germinating and growing for two years before flowering in the third year and senescing after fruiting in autumn. Perennials, such as S. perennis, overwinter as basal rosettes, persisting for multiple years before bolting and producing flowers in July-August.^[22]^[23] Asexual reproduction is rare in Swertia but occurs via rhizomes in some montane perennials, allowing clonal spread in stable habitats like those occupied by S. perennis.^[23]

Distribution and ecology

Geographic range

The genus Swertia is primarily distributed across temperate and subtropical zones of the Northern Hemisphere, encompassing parts of Asia, Europe, and North America, with notable extensions into tropical and subtropical regions of Africa and Madagascar.^[25]^[2] This broad range reflects the genus's adaptation to diverse climatic conditions, though it is absent from South America and Australia.^[26] 166 species are recognized globally, with the majority concentrated in alpine and montane environments.^[1] Centers of highest diversity occur in the Himalayan region, particularly the Sino-Himalayan area spanning China, India, and Nepal, where around 86 species have been documented, representing a significant portion of the genus's total variation.^[27] In India, approximately 36 species are recorded, primarily in the Himalayan region, while Nepal alone hosts about 29, many of which are found from subtropical elevations around 1,000 m to alpine zones up to 5,600 m across its western, central, and eastern districts.^[3]^[28]^[29] Europe supports a smaller diversity of about three species, mainly in alpine regions such as the Pyrenees, Alps, Carpathians, and other montane systems.^[30]^[31] In North America, representation includes two species: Swertia caroliniensis in the eastern United States, particularly in calcareous grasslands and savannas from southern Ontario southward to northern Alabama, and Swertia perennis in western regions.^[2]^[32]^[19] Endemism is particularly pronounced in the Sino-Himalayan region, where numerous species are restricted to localized high-altitude habitats, contributing to the area's status as a primary hotspot for the genus.^[33] Disjunct distributions are evident, such as in S. perennis, which exhibits separated populations across Eurasian mountain ranges and western North America, highlighting historical biogeographic patterns like vicariance or long-distance dispersal.^[31]^[34] No major range expansions or contractions have been widely documented to date, but post-2020 modeling studies suggest potential altitudinal migrations upward for several Swertia species in response to climate warming, particularly in Himalayan populations where suitable habitats may shift to higher elevations under future scenarios.^[35]^[36]

Habitat preferences

Swertia species predominantly inhabit montane grasslands, alpine meadows, and forest edges across temperate and tropical regions, where they thrive in open or semi-shaded environments that provide moderate sunlight and protection from extreme exposure. These plants generally avoid waterlogged soils, favoring sites with consistent but not excessive moisture to prevent root rot and support their herbaceous growth.^[3]^[37] The genus exhibits a broad climatic tolerance, spanning cool temperate zones to subtropical areas, with many species adapted to elevations between 1,000 and 4,000 meters, particularly in Himalayan hotspots. Perennial species demonstrate resilience to frost and seasonal temperature fluctuations in alpine settings, enabling persistence in high-altitude environments with short growing seasons.^[38]^[3] Soil preferences for Swertia include well-drained, acidic to neutral loams rich in humus, which facilitate nutrient absorption and root development in mountainous terrains. European species, such as Swertia perennis, often associate with calcareous substrates in fens and wet meadows, where base-rich conditions support their growth in nutrient-variable ecosystems.^[37]^[39] Symbiotic mycorrhizal associations, particularly arbuscular types, play a crucial role in nutrient uptake for Swertia, enhancing phosphorus acquisition in nutrient-poor alpine soils and aiding adaptation to stressful conditions. In open habitats, these plants compete with grasses for light and resources, often occupying niches in disturbed or successional grasslands.^[40]

Species diversity

Number of species and variation

The genus Swertia comprises approximately 166 accepted species, though estimates range from 120 to 150 due to ongoing taxonomic revisions and varying circumscriptions across regional floras.^[1]^[41] In a broader sense, including subspecies and varieties, the total exceeds 165 taxa as recognized by Plants of the World Online in 2023.^[1] Infrageneric classification divides Swertia into several sections based on morphological and distributional traits, such as Sect. Swertia (primarily European species with simple leaves and tubular corollas) and Sect. Chiraeta (Asian taxa often featuring fringed corollas and broader leaves).^[33]^[42] These groupings highlight significant intraspecific variation, including differences in corolla shape (from rotate to campanulate) and leaf size (ranging from linear to ovate), which reflect adaptations to diverse microhabitats.^[6] Genetic diversity within Swertia is elevated in polyploid species, which exhibit high heterozygosity levels due to genome duplication events, as evidenced by cytogenetic analyses in species like S. chirayita and S. angustifolia.^[43] Hybridization events have been documented in contact zones, particularly between closely related taxa in the Himalayas, contributing to further variability through introgression and allopolyploid formation.^[44] Habitat loss from deforestation, agricultural expansion, and climate-induced shifts poses a major threat to Swertia diversity, reducing populations of variant forms in alpine and montane ecosystems, especially in biodiversity hotspots like the Himalayas.^[45]^[46] Overexploitation for medicinal uses exacerbates this decline, fragmenting habitats and limiting gene flow among remaining variants.

Selected species

Swertia chirayita (Roxb. ex C.B. Clarke), commonly known as chirayita, is a prominent species native to the temperate Himalayan regions, including India, Nepal, and Bhutan, where it grows at elevations between 1,200 and 3,000 meters in grassy slopes and open forests.^[3] This annual herb reaches heights of up to 1.5 meters, featuring an erect stem that is quadrangular in the upper portion and bears opposite, sessile leaves that are elliptic and 4-6 cm long.^[47] Its bright blue flowers, arranged in a terminal panicle, bloom from September to November, contributing to its ecological role in alpine meadows.^[48] Valued for its medicinal properties in traditional systems like Ayurveda, it is harvested for treating fever, liver ailments, and digestive issues, though overexploitation has led to assessments classifying it as critically endangered using IUCN criteria, particularly in India.^[3]^[49] Swertia perennis L., the type species of the genus Swertia, is a perennial herb distributed across the circumboreal regions, including the European Alps, northern Asia, and parts of North America, thriving in wetlands such as calcareous fens, moist meadows, and streambanks at elevations up to 3,900 meters.^[50]^[51] It grows from a rhizome to 10-70 cm tall, with basal rosette leaves that are spoon-shaped and up to 22 cm long, transitioning to narrower cauline leaves.^[52] The flowers form in an open inflorescence, featuring a corolla of five bluish-purple to violet-blue lobes, each 8-14 mm long, often speckled with green or white spots and fringed nectar guides that attract pollinators like bumblebees.^[53] This species exemplifies the genus's adaptation to cold, wet environments, where it contributes to biodiversity in subalpine ecosystems.^[54] Swertia japonica (Makino) Makino is an East Asian species primarily occurring in Japan, Korea, and China, favoring mountainous forests, stream edges, and sunny grasslands at mid-elevations.^[55] This annual to biennial herb grows 5-40 cm tall from an unbranched or basally branched stem, with linear leaves 1-3.5 cm long and opposite arrangement.^[56] Its small, star-shaped flowers emerge in late summer, featuring white corolla lobes with pale purple stripes and a five-lobed calyx, adding ornamental appeal through their delicate, clustered display in shaded woodland settings.^[57] While not commercially cultivated on a large scale, its aesthetic qualities and ease in partial shade make it suitable for temperate garden ornamentation.^[55]

Phytochemistry

Major chemical constituents

Swertia species are rich in bioactive secondary metabolites, primarily secoiridoid glycosides, xanthones, alkaloids, triterpenoids, flavonoids, and phenolic compounds, which contribute to their chemical diversity.^[58] These constituents are typically extracted from aerial parts using polar solvents like methanol or ethanol, followed by chromatographic separation such as thin-layer chromatography (TLC) or high-performance liquid chromatography (HPLC) for isolation and purification.^[59] Yields of individual compounds vary by species and extraction conditions, generally ranging from 0.01 to 4% of dry weight.^[46] Secoiridoid glycosides represent one of the predominant classes, characterized by a cyclopentanopyran skeleton derived from iridoid precursors, often glycosylated at C-1. Key examples include swertiamarin, a monoterpenoid glucoside with a secoiridoid structure featuring a hemiacetal ring and glucose moiety, isolated primarily from aerial parts of species like S. chirayita via methanol extraction and ethyl acetate-methanol-water elution in chromatography, yielding up to 2-4% in S. chirayita.^[46] Gentiopicrin (gentiopicroside), another secoiridoid glycoside with a similar bicyclic structure and bitter properties, is extracted using ethanol and constitutes a major marker in S. chirayita, with concentrations around 0.1-0.5 mg/g dry weight in Nepalese species.^[59] Amarogentin, the most bitter compound in this class, shares a secoiridoid backbone but includes additional hydroxylation, isolated from root cultures or aerial extracts of S. chirayita at levels up to 0.26 mg/g.^[59] Xanthones, polyphenolic compounds with a tricyclic dibenzo-γ-pyrone core, are abundant in Swertia, often substituted with hydroxy, methoxy, or glycosyl groups. Prominent members include swertianin (1,5-dihydroxy-3-methoxyxanthone), isolated from aerial parts via hexane fractionation and chromatography in S. chirayita, and mangiferin (C-glucoside xanthone), detected across multiple species at 0.15-0.23 mg/g dry weight using methanol extraction and UV detection at 354 nm.^[60]^[59] Other notable xanthones, such as bellidifolin and methylswertianin, feature dihydroxy or dimethoxy substitutions and are separated from S. chirata extracts using silica gel chromatography.^[60] Alkaloids in Swertia are primarily indole or isoquinoline derivatives, with gentianine (a β-carboline alkaloid) being a key representative, formed via degradation of swertiamarin and detected qualitatively in all tested Nepalese species through Mayer's reagent test after methanol extraction. Gentianine is detected qualitatively in aerial parts of S. chirayita and other species, isolated by chemical conversion or direct chromatography.^[59]^[46] Triterpenoids, pentacyclic compounds based on oleanane or ursane skeletons, include oleanolic acid (3β-hydroxy-olean-12-en-28-oic acid) and ursolic acid (3β-hydroxy-urs-12-en-28-oic acid), extracted from roots or aerial parts of species like S. alata and S. angustifolia using ethanol, with quantified levels of 0.31-0.65% (3.1-6.5 mg/g) dry weight in some populations via HPTLC.^[60]^[61] These are isolated by acidification and recrystallization, contributing to the plant's overall terpenoid profile. Flavonoids, primarily C-glycosyl types, encompass isovitexin (apigenin-6-C-glucoside), a flavone with a glucose attached at C-6, isolated from S. alata aerial parts via methanol extraction. Total flavonoid content ranges from 18-26 mg quercetin equivalents per gram dry weight across species, measured by aluminum chloride colorimetric assay.^[62]^[59] Phenolic acids, such as those contributing to total polyphenols (22-67 mg gallic acid equivalents per gram), are broadly present and quantified spectrophotometrically post-methanol extraction.^[59] Species-specific compounds include swerilactones, unusual lactones with phenyl-substituted C-12 or C-13 skeletons, isolated from the whole plant of S. mileensis using ethanol extraction and chromatography, exemplifying structural novelty in this genus.^[63]

Biosynthetic pathways

The secoiridoid metabolites in Swertia species are primarily biosynthesized through the mevalonate (MVA) and methylerythritol phosphate (MEP) pathways, which generate isopentenyl diphosphate (IPP) and dimethylallyl diphosphate (DMAPP) as precursors. These condense to form geranyl diphosphate (GPP) via geranyl diphosphate synthase, followed by conversion to geraniol by geraniol synthase (GES). Subsequent hydroxylation at the 10-position by geraniol 10-hydroxylase (G10H, a cytochrome P450 enzyme) yields 10-hydroxygeraniol, which is further processed through iridodial, deoxyloganic acid, loganic acid, and secologanic acid intermediates via enzymes such as iridoid synthase (IS), 8-hydroxygeraniol oxidoreductase (8HGO), and secologanin synthase (SLS). This pathway culminates in swertiamarin formation through 7-deoxyloganetic acid 7-hydroxylase and 7-deoxyloganin glucosyltransferase (7DLGT) activities, with highest accumulation often observed in roots and flowers.^[64]^[65] Xanthone biosynthesis in Swertia integrates the shikimate pathway with acetate-malonate (polyketide) routes, often in a phenylalanine-independent manner specific to Gentianaceae. Shikimate-derived 3-hydroxybenzoic acid is activated to 3-hydroxybenzoyl-CoA by 3-hydroxybenzoate-CoA ligase (3BZL), which condenses with three malonyl-CoA units via benzophenone synthase (BPS) to produce 2,4,6-trihydroxybenzophenone. This intermediate undergoes B-ring hydroxylation by cytochrome P450 enzymes (e.g., CYP81AA subfamily) to form 2,3′,4,6-tetrahydroxybenzophenone, followed by oxidative phenol coupling and cyclization to the 1,3,5,8-tetrahydroxyxanthone core. Prenylation of this core by dimethylallyl pyrophosphate (DMAPP), catalyzed by prenyltransferases (e.g., PT8PX), introduces isoprenoid side chains in certain derivatives, enhancing structural diversity; some xanthones also incorporate secoiridoid units from secologanin for C-glycosylation.^[66]^[67] Triterpenoids in Swertia, such as oleanolic acid and rearranged swertane variants, originate from the cytosolic MVA pathway, where IPP and DMAPP form farnesyl pyrophosphate (FPP), dimerizing to presqualene diphosphate and then squalene. Squalene is epoxidized to 2,3-oxidosqualene, which undergoes cyclization by oxidosqualene cyclases (e.g., beta-amyrin synthase) to β-amyrin, the precursor to oleanane-type triterpenoids. Subsequent oxidations by cytochrome P450 enzymes (e.g., CYP716 family) introduce hydroxyl groups, yielding oleanolic acid; cycloartenol, a sterol precursor, shares early pathway steps but diverges toward phytosterols rather than these pentacyclic triterpenoids.^[68]^[69] Biosynthesis of these metabolites is tightly regulated, with upregulation under abiotic stresses such as UV radiation in alpine Swertia species like S. mussotii, where enhanced G10H and SLS expression boosts secoiridoid production as a protective response. Elicitors like methyl jasmonate (MeJA) induce cytochrome P450-mediated steps (e.g., G10H), increasing swertiamarin levels by up to twofold in treated tissues. Transcription factors such as WRKY proteins further modulate pathway genes, with tissue-specific expression (e.g., higher in roots for 7DLGT) ensuring organ-dependent accumulation.^[64]^[70] Evolutionarily, gene duplications in Gentianaceae, including Swertia, have expanded the genetic repertoire for secondary metabolite diversity, particularly through tandem and segmental duplications of WRKY transcription factors and pathway enzymes like G10H and CYP81AA. Phylotranscriptomic analyses reveal extensive duplications in the subtribe Gentianinae, correlating with the radiation of secoiridoid and xanthone profiles across high-altitude habitats, enabling adaptive chemical defenses. These events, dated to ancient polyploidizations, underscore the family's specialization in stress-responsive metabolomes.^[71]^[72]

Uses and conservation

Traditional medicine

In Ayurvedic medicine, Swertia chirayita, known as chirayata or kiratatikta, has been employed for centuries to treat fever, liver disorders such as jaundice and hepatitis, and diabetes, often through the use of decoctions or powders administered at dosages of 1–3 g per day.^[3]^[73] This herb is referenced in ancient texts like the Charaka Samhita (circa 300 BCE–200 CE), where it is recommended as a bitter tonic for reducing fever (jvaraghna) and purifying breast milk (stanyasodhana).^[74] In Tibetan medicine, species such as Swertia chirayita and S. mussotii are utilized, often under names like "zangyinchen," for addressing digestive issues, inflammation, and related conditions like hepatitis and fever, typically combined with other herbs in formulations to enhance efficacy.^[3]^[75] Preparations include soaking leaves and stems overnight to make a paste or boiling the whole plant to create an infusion, taken in small amounts such as 0.5–1 spoon daily depending on age.^[3] Other traditional systems have incorporated Swertia species empirically. In European herbalism, S. perennis (felwort) has been brewed into teas as a bitter to stimulate digestion and appetite, drawing on the gentian family's longstanding role in tonic preparations.^[76] Among Native American groups, particularly the Cherokee, S. caroliniensis (now classified as Frasera caroliniensis) was used for gastrointestinal ailments including colic, diarrhea, and stomach complaints, often as a root decoction or tonic to support overall digestive health.^[77] Common preparation methods across these traditions involve infusions, tinctures, and powders, with the plant's intense bitter taste naturally discouraging overuse and promoting moderation in consumption.^[3]

Modern research and conservation

Modern pharmacological research on Swertia species has focused on their bioactive compounds, particularly secoiridoid lactones like swerilactones, which demonstrate anti-hepatitis B virus (HBV) activity in vitro. Swerilactones A and B, isolated from Swertia mileensis, inhibit HBsAg and HBeAg secretion in HepG2.2.15 cells, with IC50 values of 3.66 mM and 3.58 mM, respectively.^[78] Similarly, antidiabetic effects have been observed through activation of the AMP-activated protein kinase (AMPK) pathway; amarogentin from Swertia chirayita enhances glucose uptake in L6 myotubes and reduces hyperglycemia in streptozotocin-induced diabetic mouse models by promoting AMPK phosphorylation, comparable to metformin.^[79] These findings build on traditional uses but emphasize mechanistic insights from animal and cell-based studies. Clinical potential remains promising yet underexplored, with S. chirayita extracts showing hepatoprotective effects in preclinical models against paracetamol-induced liver toxicity in rats, restoring enzyme levels like ALT and AST.^[80] Antioxidant assays further support therapeutic applications, as methanolic extracts of S. chirayita exhibit strong DPPH radical scavenging with IC50 values as low as 16.46 μg/mL, outperforming some synthetic standards and indicating potential for oxidative stress-related conditions.^[81] However, human trials are scarce, limited primarily to preliminary safety assessments in India, with no large-scale phase II studies confirmed as of 2025. Conservation efforts for Swertia are challenged by overharvesting in the Himalayas, driven by demand for medicinal uses, leading to population declines in wild habitats from Kashmir to Bhutan.^[82] S. chirayita, a key species, is considered critically endangered according to assessments using IUCN criteria due to habitat loss and unsustainable collection, though it lacks a global IUCN Red List assessment; reduced densities have been noted in Nepal and India since the early 2000s.^[82] Cultivation initiatives in Nepal, including farmer training in districts like Bhojpur and Panchase, have promoted ex-situ production at lower altitudes (1,200–2,000 m), yielding approximately 3.75 tonnes of dried herbage per hectare after two years.^[83] Sustainability measures include in vitro propagation protocols, which achieve high shoot multiplication rates (up to 32 microshoots per explant) from nodal or leaf segments using MS medium supplemented with BAP and GA3, enabling mass production without depleting wild stocks.^[84] Bioprospecting under the Convention on Biological Diversity's post-2020 framework emphasizes benefit-sharing and access regulations, as seen in Bhutan's initiatives for S. chirayita extracts in cosmetics, ensuring equitable use of genetic resources.^[85] Research gaps persist, including limited human clinical trials to validate efficacy and safety, with most data from in vitro or rodent models. Climate change exacerbates threats, with projections indicating 13–16% habitat loss for Himalayan Swertia species by 2050 due to rising temperatures shifting suitable alpine zones upward.^[86]^[87] In 2025, the CSIR Institute of Himalayan Bioresource Technology identified unexplored Swertia species as potential substitutes for S. chirayita to alleviate pressure on wild populations.^[88]

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https://www.researchgate.net/publication/394670271_A_new_classification_of_Gentianaceae_tribe_Gentianeae_subtribe_Swertiinae
[16]
Molecular Differentiation and Phylogeny of Swertia (Gentianaceae ...
Swertia consists of 150 species with a cosmopolitan geographic distribution and the center of species diversity in the Sino-Himalayan region.Missing: revisions | Show results with:revisions
[17]
The complete chloroplast genome and phylogentic results support ...
Swertia L. is one of the largest genera in the Gentianaceae Juss. and comprises 160 species (Ho and Liu 2015; Nampy et al. 2015; POWO 2023). Members of the ...
[18]
Swertia perennis L. | Plants of the World Online | Kew Science
Swertia. Swertia perennis L. First published in Sp. Pl.: 226 (1753). This species is accepted. The native range of this species is Europe to Caucasus, NE.
[19]
Swertia L.
### Morphology Summary for Swertia Genus
[20]
The pollination ecology of Swertia bimaculata (Gentianaceae) - PMC
Floral nectaries are closely associated with biotic pollination, and the nectar produced by corolla nectaries is generally enclosed in floral structures.
[21]
Reproductive biology of Swertia chirayita – a temperate critically ...
May 17, 2013 · Genus Swertia is represented by about 250 species (Kumar et al. 2005) and chromosome numbers for 49 species with 61 cytotypes (2n = 14, 16, 18, ...
[22]
Phylogeography of Swertia perennis in Europe based on cpDNA ...
Sep 17, 2018 · Swertia perennis(Gentianaceae) is a perennial diploid and clonal plant species that is discontinuously distributed in peat bogs in the ...
[23]
Seed biology imperative for conservation and restoration of Swertia ...
Apr 25, 2024 · Here, we studied the seed eco-physiology and regeneration potential of Swertia thomsonii-an endemic medicinal plant of western Himalaya.
[24]
Comparative and phylogenetic analyses of Swertia L ... - NIH
Swertia L., a large genus of plants in Swertiinae (Gentianaceae) with about 170 species, are mainly distributed in Asia, Africa, North America and Europe (Ho ...
[25]
[PDF] A NEW SPECIES OF SWERTIA (GENTIANACEAE) FROM THAILAND
Corolla pale purple with two dark purple bands and one green band in the basal half above the gland, the base with 2–4 linear dark blue appendages up to 0.2mm,.
[26]
Understanding reproductive biology for realization of conservation ...
The present study aims to understand reproductive biology and pollination ecology of Swertia thomsonii. •. Floral characteristics like conspicuous nectaries ...
[27]
Swertia in Nepal - Gentian Research Network
Morphology: An annual herb, 15–72 cm in length. Roots yellow, fibrous. Stems erect, terete or 4-lineolate, 0.58 cm in diameter. Internodes 0.7–4.5 cm long.Missing: fruits | Show results with:fruits
[28]
https://gentian.rutgers.edu/genera/genswernepal.htm
[29]
The molecular population structure of Swertia perennis ... - Nature
Oct 10, 2023 · The species is distributed globally across the Northern hemisphere, but also shows discontinuous distribution from Asia to North America.
[30]
Frasera caroliniensis Walter | Plants of the World Online | Kew Science
The native range of this species is SE. Canada to N. Central & E. U.S.A. It is a monocarpic perennial and grows primarily in the temperate biome.
[31]
[PDF] A taxonomic revision of Swertia L. (Gentianaceae) in South India ...
Jan 22, 2015 · The genus Swertia was originally established by Linnaeus (1753), who included five species in it. In the subsequent decades, a number of genera ...Missing: confusion | Show results with:confusion
[32]
[PDF] Phylogeography of Swertia perennis in Europe based on cpDNA ...
Sep 17, 2018 · The species is distributed globally across the Northern hemisphere, but also shows discontinuous distribution from Asia to North America. It ...
[33]
Will They Still Be Together? Distribution Modeling of Six Co ... - MDPI
The genus Swertia L. belongs to the family Gentianaceae, it comprises about 150 species, mainly distributed in Asia and Africa with a few species in North ...
[34]
a Himalayan endemic medicinally important plant under climate ...
Dec 20, 2022 · Our study revealed that under current climatic scenarios, the suitable habitats for the species occur across the western Himalayan region.
[35]
Swertia American Columbo PFAF Plant Database
... soils and prefers well-drained soil. Suitable pH: mildly acid and neutral soils. It can grow in semi-shade (light woodland) or no shade. It prefers moist soil.Missing: preferences | Show results with:preferences
[36]
Swertia - an overview | ScienceDirect Topics
Swertia is a diverse genus of herbs belonging to the family Gentianaceae and comprising roughly 170 species distributed throughout the world.
[37]
(PDF) Isozyme variability of the wetland specialist Swertia perennis ...
Aug 5, 2025 · ... habitat of many. specialist species, and calcareous fens harbor nearly 50% of. all endangered plant species in Switzerland (Landolt, 1991).
[38]
The Role of Arbuscular Mycorrhiza in the Growth and Development ...
Arbuscular mycorrhiza (AM) is an ancient symbiosis between land plants and those fungi from the Phylum Glomeromycota. It is estimated to occur in a large ...
[39]
The complete chloroplast genome and phylogentic results support ...
Apr 24, 2024 · Swertia L. is one of the largest genera in the Gentianaceae Juss. and comprises 160 species (Ho and Liu 2015; Nampy et al. 2015; POWO 2023).
[40]
A taxonomic revision of Swertia L. (Gentianaceae) in South India ...
The genus Swertia in South India is revisited based on field and herbarium studies. A brief history of the genus, detailed descriptions, photo plates, ...
[41]
Cytogenetic and DNA fingerprinting analysis in three species of ...
May 7, 2015 · Thus, the present study reveals DNA markers that can be utilized for identification of genetic diversity of Swertia species. Genetic ...
[42]
Introgression of Swertia mussotii gene into Bupleurum ...
The hybridity of the calli was confirmed by a combination of isozyme, RAPD and chromosomal analysis. The hybrid calli genomes were predominantly B.Missing: polyploidy | Show results with:polyploidy
[43]
a Himalayan endemic medicinally important plant under climate ...
Dec 20, 2022 · In the current era of the anthropocene, climate change is one of the main determinants of species redistribution and biodiversity loss.
[44]
A Review of Swertia chirayita (Gentianaceae) as a Traditional ...
Swertia, a genus in the family Gentianaceae include a large group of annual and perennial herbs, representing approximately 135 species. Swertia species are ...Missing: Linnaeus confusion
[45]
Swertia chirayita - eFlora of India
Dec 23, 2024 · Tall herb reaching 1.5 m, obscurely angled;cauline leaves subsessile, elliptic, 4-6 cm long, 5-nerved, lower much longer and often petiolate.
[46]
Swertia chirayita - Flowers of India
Oct 24, 2012 · Chirayita has an erect, about 2–3 ft long stem, the middle portion is round, while the upper is four-angled, with a prominent decurrent line at ...Missing: height | Show results with:height
[47]
Morphological Diversity and Nomenclature of Swertia chirayita ...
Apr 2, 2016 · Swertia chirayita is a highly endangered and very well known medicinal plant enlisted in the IUCN list of endangered plants in the world.
[48]
Swertia - Wikispecies - Wikimedia
Jul 12, 2024 · Name. edit. Swertia L., Sp. Pl. 1: 226. (1753). Type species: Swertia perennis L., Sp. Pl. 1: 226. (1753). Lectotype, designated by Hitchcock ...
[49]
FNA: Swertia perennis - Northwest Wildflowers
Swertia perennis. Phenology, Flowering summer–early fall. Habitat, Wet meadows, thickets, bogs, stream banks. Elevation, 0–3900 m. [0–12800 ft.] Distribution.
[50]
(species) swertia perennis - Montana Field Guide
Swertia is a perennial herb with erect, glabrous stems 10-13 cm tall from a rhizome. Basal leaves 4-22 cm long with long petioles and spoon-shaped to rounded ...
[51]
Swertia perennis - Burke Herbarium Image Collection
Habitat: Streambanks, wet meadows and other moist places, mid- to high elevations in the mountains. Flowers: July-August. Origin: Native. Growth Duration ...
[52]
Swertia perennis (Felwort) | Native Plants of North America
Mar 28, 2025 · Several erect stems, with most leaves at base, and star-like, pale bluish-purple flowers with greenish or white spots in an open, branched, but ...
[53]
Frasera caroliniensis - Species Page - NYFA - New York Flora Atlas
Annual: Plants that perform their entire lifecycle within a single growing season. All roots, stems, and leaves die at the end of the growing season. Over ...
[54]
Frasera caroliniensis: The Anticipation of an Impending Bloom
May 28, 2021 · It will grow as a low mound for many years, sometimes as long as 30, before flowering! The 26 Native Plant Garden specimens were planted in 2011 ...Missing: Swertia habitat annual white
[55]
American Columbo - Flora of Pennsylvania - PAEnflowered
Its flowers have four large sepals that branch out from the central ovary and four white to light green petals, each with purple spots and a conspicuous, ...Missing: annual dots
[56]
Frasera caroliniensis (American columbo) | Native Plants of North ...
Jan 23, 2023 · Found in calcareous grasslands and savannah over much of east-central North America, but not common, Frasera caroliniensis is listed as a species of special ...
[57]
Swertia japonica - Useful Temperate Plants
Swertia japonica is an annual to biennial plantwith stems that are unbranched or branched from the base; it can grow around 5 - 40cm tall.
[58]
Swertia japonica PFAF Plant Database
The species is hermaphrodite (has both male and female organs). Suitable for: light (sandy), medium (loamy) and heavy (clay) soils. Suitable pH: mildly acid, ...
[59]
Swertia japonica | Flower Database
Swertia japonica, or Senburi, is a biennial plant native to Japan, Korea, and China. It has star-shaped white flowers with purple streaks, and is used as a ...
[60]
The American Journal of Chinese Medicine
### Summary of Phytochemical Content of Swertia Species
[61]
Phytochemical investigation of crude methanol extracts of different ...
Dec 26, 2015 · The genus Swertia is reported to contain potent bitter compounds like iridoids, xanthones and c-glucoflavones that are known to heal many human ...
[62]
[PDF] Rapid HPLC-PDA Estimation of Swertiamerin in Extracts of ...
The yields of the cold (room temperature) methanol and water extraction of the aerial parts were found to be 45.8% and 41.5% and that of hot methanol and water ...<|separator|>
[63]
Chemical constituents and biological importance of Swertia: a review
Swertia species are known to contain various active components including antioxidant xanthones, seco-irioid glycosides, flavonoids, alkaloids, and phenolics 31, ...
[64]
(PDF) Determination of Ursolic Acid a Biomarker in Different Swertia ...
Aug 7, 2025 · A simple highly precise method has been established for the determination of ursolic acid in three species of Swertia viz. S. angustifolia, S.
[65]
EVALUATION OF TOTAL PHENOLIC, FLAVONOID, INVITRO ...
Besides these compounds, isovitexin (Ia) has been isolated from the plant and identified. At the same time, flavonoid constituents were examined with five kinds ...
[66]
Swerilactones E–G, three unusual lactones from Swertia mileensis
Swerilactones E (1), F (2), and G (3), three unusual lactones with a phenyl group, were isolated from the traditional Chinese herb of Swertia mileensis.
[67]
Deep sequencing and transcriptome analyses to identify genes ...
Feb 22, 2017 · To better understand the secoiridoid biosynthesis pathway, we generated transcriptome sequences from the root, leaf, stem, and flower tissues, ...
[68]
Contents of therapeutic metabolites in Swertia chirayita correlate ...
Swertiamarin and amarogentin take the classical MVA/MEP route of terpene biosynthesis followed by the secoiridoid pathway (Fig. 1). These secoiridoids ...
[69]
Xanthone Biosynthetic Pathway in Plants: A Review - Frontiers
The pathway initially involves the shikimate pathway, either through L-phenylalanine-dependent or -independent pathway, that later forms an intermediate ...Abstract · Introduction · Biosynthesis of Xanthone Core... · Conclusion
[70]
Structure, activity and biosynthesis of xanthones in genus Swertia
Oct 25, 2025 · Biosynthetic pathways are complex, involving multiple enzymes and intermediates. Among them, the study of some key steps and enzymes provides ...
[71]
Triterpenoid Biosynthesis and Engineering in Plants - Frontiers
Triterpenoid saponins are a diverse group of natural products in plants and are considered defensive compounds against pathogenic microbes and herbivores.Missing: Swertia | Show results with:Swertia
[72]
Two New Chiratane-Type Triterpenoids from Swertia kouitchensis
Two rare new chiratane-type triterpenoids, kouitchenoids A and B (1, 2), together with oleanolic acid (3) and ursolic acid (4), were isolated from ethanol ...Missing: major | Show results with:major
[73]
The role of UV-B radiation in modulating secondary metabolite ...
Furthermore, UV-B radiation induces gene regulation in secondary metabolism, with most genes being upregulated. UV-B interacts with other stress factors, e.g. ...Introduction · Uvr8 Signaling Regulates... · Uv-B Regulation Of Terpenoid...
[74]
Genomic characterization of WRKY transcription factors related to ...
Jan 23, 2024 · In simple terms, the secoiridoid biosynthesis pathway in plants can be divided into three main parts: the initial MEP and MVA pathways and the ...
[75]
Phylotranscriptomics reveals extensive gene duplication in the ...
In this study, we aim to reconstruct the genome‐scale phylogeny and identify large‐scale gene duplication events for the subtribe Gentianinae (Gentianaceae), ...
[76]
Kiratatikta - Swertia chirata - Benefits, Dose, Usage, Side Effects
Jan 8, 2014 · It destroys diseases because of its bitter taste. Bhunimb: Bhuva: Nimb: Iva: It is a small neem plant or herb. Kandathiktha: Thiktha Kanda ...
[77]
[PDF] Local Knowledge on the Use of Swertia chirayita as Traditional ...
Dec 7, 2015 · and Tibetan (Sharma et al. 2008). The whole plant of S. chirayita is used in traditional medicine to treat fever, ma- laria, cough, cold ...
[78]
Natural Medicines Used in the Traditional Tibetan Medical System ...
Jan 30, 2018 · It has been commonly used for the treatment of hepatitis, acute icterohepatitis, cholecystitis, epidemic fever, headache, and blood diseases in ...
[79]
Gentian Uses, Benefits & Side Effects - Drugs.com
The gentians have been used for centuries as bitters to stimulate the appetite, improve digestion, and to treat a variety of GI complaints.
[80]
Frasera caroliniensis Walt. - Native American Ethnobotany Database
Frasera caroliniensis Walt. Common names: American Columbo Species details (USDA): USDA FRCA2. Documented uses. 6 uses documented
[81]
Swerilactones A and B, anti-HBV new lactones from a ... - PubMed
Swerilactones A and B, anti-HBV new lactones from a tradtional Chinese herb: Swertia mileensis as a treatment for viral hepatitis · Abstract · Publication types.
[82]
Amarogentin, a secoiridoid glycoside, activates AMP
Glucose uptake in L6 myotubes and endothelial inflammation were assessed to determine the functional effects of AMPK activation. ... animal models, and human ...
[83]
Hepatoprotective activity of Andrographis paniculata and Swertia ...
The present study revealed that the extracts of A. paniculata or S. chirayita offered protection against hepatotoxicity induced by paracetamol.Missing: phase 2023
[84]
[PDF] A Comparative Evaluation of Swertia Species as Potential ...
Jun 1, 2014 · However, only methanolic extract of S. chirata and S. lurida exhibited maximum in vitro antioxidant activity in DPPH assay shown IC50 16.46 and ...
[85]
Ex-situ cultivation at lower altitude and evaluation of Swertia ...
S. chirayita is categorized “critically endangered” by the International Union for Conservation of Nature (IUCN) (Joshi and Dhawan, 2005). Depleting population ...
[86]
Production from both wild harvest and cultivation: The cross-border ...
Oct 28, 2018 · Swertia chirayita is the most widely traded species in a genus of 150 species, many of which are used in traditional medicine.
[87]
Swertia Chirayita: Medicinal Plant - The Wonder Nepal
May 23, 2025 · For instance, in Bhojpur, Nepal, over 70 farmers have adopted commercial farming of Swertia chirayita, cultivating it at altitudes between 1, ...
[88]
Efficient In Vitro Propagation Protocol of Swertia chirayita (Roxb. ex ...
Apr 10, 2018 · The results of the present study showed that the leaf explants were good source for in vitro propagation of S. chirayita via callus induction ...
[89]
Acts, rules & policies - National Biodiversity Centre
Promoting Bioprospecting ... Launch of SUBLIMAGE L'EXTRAIT using Bhutanese Swertia chirayita – From Lauri Gewog, Samdrup Jongkhar to Chanel Parfums Beaute, France ...
[90]
Medicinal plants in peril due to climate change in the Himalaya
A total of 13–16% of medicinal plant species currently found in the region are likely to lose their existing potential habitats by 2050 and 2070.
[91]
Climate change‐induced distributional change of medicinal and ...
Aug 15, 2022 · This study identifies the potential distribution of 29 species of MAPs in Nepal under current and future climate using an ensemble modeling and hotspot ...