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Atropa
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Solanales
Family: Solanaceae
Subfamily: Solanoideae
Tribe: Hyoscyameae
Genus: Atropa
L. (1753)[1]
Species[2]

6, see text

Synonyms[2]
  • Belladona Mill. (1754), nom. superfl.
  • Pauia Deb & Ratna Dutta (1965)

Atropa is a genus of flowering plants in the nightshade family, Solanaceae: tall, calcicole, herbaceous perennials (rhizomatous hemicryptophytes), bearing large leaves and glossy berries particularly dangerous to children, due to their combination of an attractive, cherry-like appearance with a high toxicity.[3] Atropa species favour temperate climates and alkaline soils, often growing in light shade in woodland environments associated with limestone hills and mountains.[4][5] Their seeds can remain viable in the soil for long periods, germinating when the soil of sites in which plants once grew (but from which plants have long been absent) is disturbed by human activity[6] or by natural causes, e.g. the windthrow of trees (a property shared by the seeds of other Solanaceae in tribe Hyoscyameae e.g. those of Hyoscyamus spp., the henbanes).[7] The best-known member of the genus Atropa is deadly nightshade (A. belladonna) – the poisonous plant par excellence in the minds of many.[8] The pharmacologically active ingredients of Atropa species include atropine, scopolamine, and hyoscyamine, all tropane alkaloids having anticholinergic, deliriant, antispasmodic and mydriatic properties.[citation needed] The genus is named for Άτροπος (Atropos) – lit. 'she who may not be turned (aside)' – one of the Three Fates and cutter of the thread of life / bringer of death – in reference to the extreme toxicity of A.belladonna and its fellow species – of which four others are currently accepted.[9][10]

In some older classifications, the mandrake (Mandragora officinarum) has been placed in the genus Atropa, under the binomial Atropa mandragora.

Species

[edit]

Six species and one natural hybrid are accepted.[2]

The genus Atropa is currently under review, so changes in nomenclature are likely, once said review is complete. It will be seen from the above that there is an overlap in the respective distributions of A. acuminata, A. komarovii and A. pallidiflora in the lush Hyrcanian forests of Northern Iran, and it is possible that some or all of these species may yet be subsumed in the concept Atropa belladonna. A. belladonna itself (including its variety caucasica) is also present in the Hyrcanian forests and vol. 100 of Flora Iranica includes a useful key with which to distinguish the four species occurring in northern Iran. Data on A. pallidiflora and A. acuminata Royle ex Miers are neither abundant nor readily accessible on the Internet at present. The reported presence of an Atropa species in Mongolia is intriguing, given that country's relative remoteness from Kashmir and its (Kashmir's) well-attested population of Atropa acuminata Royle ex Lindl. The unequivocal presence of Atropa in the Eastern Himalaya would go at least some way to bridging the gap between Kashmiri and Mongolian populations of this genus. Some light might be cast upon this problem by the gaining of better knowledge concerning the rare and poorly-known species A. indobelladonna, found in Arunachal Pradesh and adjoining areas of Assam. This was first described in 1961 under the name Pauia belladonna, as the sole species of the monotypic genus Pauia, the specific name belladonna being bestowed by authors Deb and Dutta because of its partial similarity to Indian belladonna (Atropa acuminata Royle ex Lindl).[13] In 2020 It was placed in genus Atropa, on the strength of evidence scant, to say the least, and somewhat at variance with the original anatomical line drawings of the species. No photographic images of the species are currently viewable on the internet.[14]

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Atropa

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from Grokipedia
Atropa is a small genus of flowering plants in the nightshade family, Solanaceae, comprising approximately four to six accepted species of rhizomatous perennial herbs native to Europe, western and central Asia, and North Africa. These plants are characterized by branching stems, alternate leaves (sometimes in pairs), solitary axillary flowers that are five-merous with campanulate calyces and corollas in shades of purple or yellow, and globose fleshy berries. The chromosome number is typically x = 36. The most notable species is Atropa bella-donna L., commonly known as deadly nightshade or belladonna, which is a multi-stemmed herbaceous growing 1–2 meters tall with dark green ovate leaves, dull purple bell-shaped flowers from to , and shiny black berries. Native to soils in and the Mediterranean region, it has been introduced and naturalized in parts of , such as and the , often in disturbed areas like roadsides and waste places. Other accepted species include A. acuminata Royle ex Lindl., found in eastern to ; A. baetica Willk., endemic to central and southern and ; A. pallidiflora Schönb.-Tem., native to northern to the ; A. komarovii Blin. & Shalyt, occurring in northeastern to southern ; and A. indobelladonna Karthik. & V.S.Kumar, native to , . Members of the Atropa are renowned for their toxicity due to the presence of alkaloids such as atropine, , and , which act as anticholinergics and are concentrated in all plant parts, particularly the berries and roots. Ingestion can cause severe symptoms including dilated pupils, rapid heartbeat, , hallucinations, fever, and potentially fatal , with as little as 600 mg of dried plant material being lethal to adults. Despite their poisonous nature, these alkaloids have significant medicinal value; atropine, derived primarily from A. belladonna, is used in to dilate pupils, as an for certain poisonings, and in treatments for conditions like and exposure. Historically, extracts were employed in folk for pain relief and as a cosmetic to enlarge pupils, though their use requires caution due to the narrow . The genus name Atropa derives from , one of the three in who cuts the thread of life, alluding to the deadly potential of its species. While A. belladonna has cultural associations with and in , the plants generally prefer well-drained, calcareous soils in full sun to partial shade and are not recommended for cultivation in home gardens due to their .

Taxonomy

Etymology

The genus name Atropa derives from , one of the three (Fates) in , who was responsible for severing the thread of life with her shears, symbolizing an inexorable end. This name was selected by to reflect the highly toxic and often fatal properties of the plants in the genus, evoking the inescapable doom associated with their poisoning. Linnaeus formally established the genus Atropa in his seminal work Species Plantarum (volume 1, page 181), published in 1753, where he introduced binomial nomenclature for plants and highlighted the deadly nature of these species through the mythological allusion. The choice underscores the historical recognition of the genus's potent alkaloids, which can lead to rapid and irreversible physiological effects.

Classification

Atropa is a genus of flowering plants classified within the family Solanaceae, order Solanales, class Magnoliopsida, phylum Tracheophyta, and kingdom Plantae. Within Solanaceae, it belongs to the subfamily Solanoideae and the tribe Hyoscyameae. Phylogenetic analyses place Atropa in close relation to genera such as Hyoscyamus and Anisodus within the tribe Hyoscyameae, with molecular studies based on chloroplast DNA and nuclear markers indicating divergence in temperate regions of Eurasia. This divergence is supported by evidence of ancient polyploidy events and shared retroposon insertions that distinguish Atropa as an early-branching lineage in the tribe. The genus Atropa was first described by in his 1753 work , where he established Atropa belladonna as the . Taxonomic revisions in the 19th and 20th centuries refined species boundaries, with notable additions including Atropa baetica described in 1852 and Atropa pallidiflora in 1972, addressing morphological variations across Eurasian distributions. As of 2025, recognizes six accepted species in the genus, though debates persist regarding the status of certain hybrids and potential synonyms. Historical misclassifications have included the placement of some Atropa species in separate genera, such as early synonymy of Atropa acaulis with , later corrected through morphological and molecular reassessments. One ongoing point of contention involves the hybrid Atropa × martiana (a cross between A. baetica and A. belladonna), which is accepted as a distinct nothospecies in regions of overlap in and but raises questions about natural hybridization and species delimitation.

Species

The genus Atropa includes six accepted , all toxic native to temperate regions of and , distinguished primarily by variations in flower color, fruit morphology, and geographic isolation.
  • Atropa acuminata Royle ex Lindl., known as Indian belladonna, is a branched up to 1.6 m tall with fistular stems, ovate-lanceolate leaves 8–17 cm long, and solitary nodding flowers.
  • Atropa baetica Willk., or Andalusian belladonna, is a rhizomatous reaching 1.7 m with erect glabrous stems, pale greenish-yellow flowers veined in , and green berries turning black; it is considered threatened due to its rarity.
  • Atropa bella-donna L., commonly called deadly nightshade, features dull bell-shaped flowers and glossy black berries on plants up to 2 m tall.
  • Atropa indobelladonna Karthik. & V.S. Kumar, a recently described from 2020, is a similar to A. acuminata but with more acuminate leaves and flowers adapted to high-altitude habitats.
  • Atropa komarovii Blin. & Shalyt. is a taller up to 1.8 m with broad leaves, greenish-yellow flowers veined , and soft berries.
  • Atropa pallidiflora Schönb.-Tem. is a shrubby 20–90 cm tall with pale flowers and berries containing alkaloids.
One natural hybrid is recognized: Atropa × martiana Font Quer, resulting from A. baetica × A. belladonna. All species exhibit shared toxicity from tropane alkaloids like atropine and scopolamine.

Description

Morphology

Atropa species are herbaceous perennials classified as hemicryptophytes, characterized by their rhizomatous root systems that enable vegetative persistence and regeneration. These plants typically grow to heights of 1-2 meters, forming robust, erect habits suited to their native woodland and scrub environments, though heights vary slightly among species (e.g., up to 1.6 m in A. acuminata). The roots are thick, fleshy, and branched, often whitish in color, providing anchorage and nutrient storage. Stems emerge from the rhizomes as erect and branching structures, reaching up to several branches per , with surfaces that range from glabrous to sparsely pubescent or glandular-hairy. Leaves are arranged alternately along the stems, though subopposite pairs (geminate) occur particularly on upper portions, measuring 10-30 cm in length and featuring ovate to lanceolate shapes with entire margins, acute apices, and prominent veins on the undersurface; they are borne on short petioles and exhibit a dull dark green coloration, with minor variations in size across species. Flowers develop solitarily in axils, pendulous and nodding, with campanulate corollas 2-3 cm long that are five-lobed and to reflexed at the mouth, colored in (in A. belladonna) or yellow to greenish-yellow (in other ), often with veins or tinges. The persistent calyx is accrescent and campanulate, featuring five acute to acuminate lobes that widen and in . Fruits are juicy, globose berries, 1-2 cm in diameter, glossy and ranging from black to across , each containing numerous small . Seeds are disc-shaped to globose-reniform, with high viability that allows persistence in soil seed banks for 3-5 years, facilitating opportunistic .

Phytochemistry

Atropa species, particularly A. belladonna, are renowned for their rich content of alkaloids, a class of bioactive nitrogenous compounds that define the genus's pharmacological significance. The primary alkaloids include (the naturally occurring L-form), atropine (the racemic dl-), and (also known as hyoscine). These alkaloids accumulate in varying concentrations across plant parts, with the highest levels typically found in and fruits; for instance, atropine reaches approximately 10 mg/g dry weight in the main root and 3 mg/g in leaves, while concentrations are generally lower but notable in reproductive tissues. In addition to tropane alkaloids, Atropa plants contain other secondary metabolites, including solanaceous steroidal glycosides such as spirostane-type compounds (e.g., atroposides A–H), which contribute to the plant's chemical diversity. , including derivatives of , , , and scopolin, are present primarily in leaves and provide properties. Withanolides, a group of steroidal lactones characteristic of the family, have also been identified in Atropa, though in lower abundances compared to related genera like Withania. The biosynthesis of tropane alkaloids in Atropa proceeds from the amino acid , which is decarboxylated to by , a rate-limiting step. is then methylated to N-methylputrescine, which cyclizes to form the 4-N-methylaminobutanal intermediate, ultimately leading to —the bicyclic precursor—through a non-enzymatic condensation involving an unusual type III . From , reduction yields or pseudotropine, which are further acylated to produce and, via epoxidation and hydrolysis, . Alkaloid profiles in Atropa exhibit significant variation, influenced by developmental stage, environmental factors, and . Scopolamine concentrations are notably higher in fruits and sepals compared to vegetative parts, often exceeding 1 mg/g dry weight in fruit sepals. Genetic factors, including differential expression of genes like hyoscyamine 6β-hydroxylase (h6h) and tropinone reductases (TR1 and TR2), play a key role in determining hyoscyamine-to-scopolamine ratios, as revealed through genomic analyses of Atropa species.

Distribution and habitat

Geographic range

The genus Atropa is native to temperate regions of and , with species distributed across a wide latitudinal range from to the . In , A. belladonna occupies much of the continent, extending from and eastward to the and , while A. baetica is more restricted to disjunct populations in the central and southern . These European ranges connect to North African extensions of A. belladonna in and , forming a continuous Mediterranean distribution. In , the genus shows greater fragmentation, with A. acuminata ranging from eastern through , , and the into northern , and A. indobelladonna limited to in northeastern . Further west, A. pallidiflora occurs in the of northern , and A. komarovii in northeastern and southern , reaching into . These Asian locales highlight disjunct populations adapted to montane and woodland edges. Beyond its native range, A. belladonna has been introduced and naturalized in parts of , particularly in the and southeastern , as well as in , where it persists in disturbed areas. Biogeographically, centers of diversity for Atropa lie in the Mediterranean-Turanian region and the Himalayan-Qinghai-Tibet Plateau area, reflecting post-glacial migrations from southern refugia that allowed northward and eastward recolonization following the .

Habitat preferences

Species of the genus Atropa are calcicole plants, thriving in calcium-rich, alkaline soils with a pH typically ranging from 7 to 8. They prefer well-drained loamy soils that retain moisture without becoming waterlogged, such as light, permeable chalky substrates often found on south-west facing slopes. These conditions support optimal growth and alkaloid production, particularly in A. belladonna, where calcareous soils enhance medicinal compound concentrations. The genus favors temperate climates characterized by cool winters and mild to warm summers, aligning with Mediterranean and Euro-Siberian influences. Plants exhibit tolerance to light shade, growing effectively in partial sunlight or dappled woodland conditions, though full sun exposure maximizes vigor and secondary metabolite yields. Germination and establishment occur in late winter to early spring under fluctuating temperatures, with optimal regimes around 32°C day/18°C night for species like A. baetica and A. belladonna. Atropa species occupy specific microhabitats including edges, scrublands, rocky slopes, and karsts, often in disturbed areas such as path banks and clearings. They are commonly associated with (Quercus) and (Fagus) s, colonizing understories in mixed upland woodlands of Quercus faginea, Quercus ilex, Quercus pubescens, and Fagus sylvatica. Elevations range from sea level to 2000 m, with A. baetica favoring montane sites above 900 m in Iberian pine (Pinus nigra) stands. Adaptations to these environments include a robust system enabling by accessing deeper soil moisture reserves. In shaded microhabitats, plants display as a shade avoidance mechanism, promoting stem elongation to compete for light while maintaining . Seed dormancy is physiologically nondeep, alleviated by cold stratification or , facilitating persistence in short-lived soil seed banks (3–5 years) within variable clearings.

Ecology

Reproduction

Atropa species exhibit hermaphroditic flowers that bloom during the summer, typically from June to August in temperate regions. These flowers are self-compatible, enabling autogamous pollination within the same flower due to their pendulous structure, which facilitates pollen transfer as anthers dehisce. However, outcrossing is predominantly promoted through entomophily, with primary pollinators including bees (Apis spp.) and bumblebees (Bombus spp.), which vibrate the flowers to release pollen via buzz pollination. Pollination is supported by floral structures adapted for visitation, though rewards are minimal and contain toxic alkaloids such as and , which can deter prolonged feeding but do not prevent effective transfer by pollinators like bumblebees. These alkaloids in and may select for tolerant pollinators, ensuring despite the plant's toxicity. occasionally visit, contributing to secondary . After successful , the tetralocular ovaries develop into berries that ripen in autumn, generally from to , containing 25–35 per fruit in Atropa belladonna. occurs primarily via endozoochory, with birds consuming the attractive, glossy black berries and excreting viable , despite the fruits' high content; certain avian exhibit tolerance to these toxins, facilitating long-distance dispersal. As perennials, Atropa persist through vegetative via rhizomes, enabling clonal expansion and survival in suitable habitats. demonstrate conditional physiological , requiring alternating temperatures (e.g., 32/18°C) for , which can be erratic and take 1–6 months; viability persists for up to 5 years in banks, supporting population persistence.

Interactions

Atropa species experience limited herbivory primarily due to the presence of toxic alkaloids, such as atropine and , which act as potent chemical defenses against most mammalian herbivores. These compounds induce narcosis, , or in generalist feeders like humans and many domestic animals, resulting in low browsing pressure from mammals in natural habitats. However, some mammals, including and rabbits, appear tolerant and can graze on the foliage without significant adverse effects, possibly due to physiological adaptations or lower concentrations in certain plant parts. Among , herbivory is similarly reduced, but specialized feeders—such as certain lepidopteran larvae or coleopterans adapted to -rich hosts—can tolerate and consume Atropa tissues, exerting selective pressure on production. These interactions highlight the alkaloids' role in balancing defense against diverse herbivores while allowing limited consumption by tolerant species. In terms of symbioses, Atropa plants commonly form mutualistic associations with arbuscular mycorrhizal fungi (AMF) from phyla such as Glomeromycota, which colonize cortical cells to form arbuscules and enhance acquisition. This is particularly beneficial in -poor soils, where AMF extend the system's absorptive surface, improving uptake of immobile nutrients like and , thereby supporting growth and survival in oligotrophic understories. Studies on , including Atropa, indicate high colonization rates (often exceeding 50% of root length) by AMF genera like Glomus and Rhizophagus, which also contribute to and overall vigor in suboptimal conditions. The reciprocal exchange involves the providing carbohydrates to the fungi, fostering a stable partnership that aids Atropa's persistence in shaded, infertile habitats. Competition among Atropa and co-occurring understory plants is influenced by its and chemical interference via . As a thriving in partial shade beneath woodlands, Atropa competes effectively with other low-light adapted for , , and , often dominating in disturbed edges. Its tropane alkaloids, leached from , leaves, or decaying tissues, exert allelopathic effects by inhibiting seed germination and seedling growth in neighboring through disruption of cellular processes like activity and integrity. For example, atropine has been demonstrated to suppress mitotic divisions and root elongation in test , reducing competitive pressure from sympatric and facilitating Atropa's establishment. These interactions, mediated by the same alkaloids central to its , underscore Atropa's strategy for niche occupancy in diverse ecosystems. Pathogen interactions pose significant threats to Atropa, especially in cultivated settings, with susceptibility to both fungal and viral agents. The soilborne fungus Verticillium dahliae causes , invading vascular tissues to induce wilting, yellowing, and eventual plant death; Atropa belladonna serves as a wild host, harboring the and contributing to its persistence in Solanaceae-dominated areas. Symptoms typically emerge under stress, with microsclerotia surviving in soil for years and exacerbating disease in nutrient-poor or waterlogged conditions. Virally, Atropa is prone to infections by Belladonna mosaic virus (BeMV) and mosaic virus (ToMV), which manifest as leaf mottling, chlorotic spots, ringspots, and necrotic lesions, severely impacting and growth. These viruses, often vectored by mechanical means or in cultivation, can lead to systemic spread and reduced vigor, with BeMV specifically noted for causing mottle patterns in natural infections. Such susceptibilities highlight vulnerabilities despite chemical defenses, particularly when grown for extraction.

Uses

Medicinal

Atropa species, particularly Atropa belladonna, have been utilized in for centuries, with ancient civilizations employing plant extracts to alleviate pain and treat respiratory conditions such as . In and , belladonna was used as a and , often in the form of ointments or ingested preparations to manage labor pains and inflammatory disorders. By the , it appeared in European herbal remedies for conditions like and as an for gastrointestinal issues. The key medicinal alkaloids atropine and were first isolated from Atropa belladonna in 1833 by German pharmacists Philipp L. Geiger, Rudolf Brandes, and later confirmed by Heinrich F. G. Mein, enabling more precise pharmaceutical applications. This isolation marked a pivotal advancement, shifting from crude herbal uses to standardized extractions for therapeutic purposes. In modern , atropine, derived from Atropa, serves as a primary agent for dilation in ophthalmological examinations and procedures, administered topically to paralyze the iris and ciliary muscles. It is also used intravenously to treat by blocking parasympathetic activity on the heart, with an initial dose of 1 mg IV repeated every 3-5 minutes up to a total of 3 mg in adults (as of 2020 AHA guidelines). Scopolamine, another Atropa alkaloid, is employed transdermally or orally to prevent and intravenously to reduce during , often at doses of 0.3 to 0.65 mg. Derivatives of these alkaloids feature in combination drugs like , which includes atropine, scopolamine, and for and other antispasmodic needs. Controlled cultivation of Atropa plants and systems, such as hairy root cultures induced by Agrobacterium rhizogenes, has been developed to sustainably produce these alkaloids, bypassing wild harvesting and ensuring consistent yields for pharmaceutical extraction. Preparations include tinctures, where belladonna leaf extract is diluted in alcohol at 0.3 mL doses for oral use, and injectables like atropine sulfate solution (0.4–1 mg/mL) for rapid systemic delivery. These compounds are regulated as prescription medications in many countries, with certain Atropa extracts classified under schedules, such as Schedule II in the United States when combined with opioids, to prevent misuse.

Other

Atropa species, particularly A. belladonna, have been cultivated ornamentally for their striking purple bell-shaped flowers and glossy black berries, adding dramatic flair to gardens despite their high requiring careful handling and placement away from children and pets. Modern ornamental use continues in specialized "poison gardens," such as the one at in , where it is showcased for educational purposes with prominent toxicity warnings to prevent accidental . Historically, Atropa plants hold significant cultural associations in , notably with ; A. belladonna was a key ingredient in "flying ointments" from the onward, applied transdermally to induce hallucinatory experiences interpreted as enabling witches to "fly" to gatherings, often combined with other sources like henbane. Folk uses extended to rituals in , such as strengthening wine for ceremonial purposes in the or as a in and warfare, while in parts of Minor, similar intoxicating applications appeared in shamanic practices, though less documented. Industrially, Atropa species serve a minor but targeted role as sources of tropane alkaloids like and for pharmaceutical research and production, with optimized drying and storage methods preserving alkaloid yields up to 21 mg/g in leaves for commercial extraction. Due to their potent , Atropa have no significant or broad agricultural applications, instead posing risks of contamination in crops like cereals and , as evidenced by regulatory assessments of tropane alkaloids as undesirable substances. Cultivation of Atropa typically involves from sown fresh in a or , where of stored can take 1–6 months at 10°C, followed by out seedlings into pots for overwintering before in late spring. Cuttings from mature plants can also be rooted in controlled environments to ensure establishment, though the process demands moist, well-drained soil and protection from to support the growth habit. All species of Atropa are highly toxic due to the presence of tropane alkaloids, primarily atropine, , and , which function as s. These compounds are found throughout the plant but are most concentrated in the berries, roots, and leaves. of Atropa can lead to , with symptoms including dry mouth, dilated pupils, , , , , , , confusion, hallucinations, , and agitation. Severe cases may progress to seizures, , and , potentially resulting in death. The toxicity is similar across species, though most documented cases involve A. belladonna. For adults, as few as 10 berries or one leaf of A. belladonna can be fatal, while 2–3 berries may be lethal to children. The minimum of atropine itself is approximately 10 mg in adults, though this varies with individual factors such as age, health, and concurrent exposures. Other species like A. acuminata, A. baetica, A. pallidiflora, and A. komarovii contain comparable profiles and pose equivalent risks.

Conservation

The conservation status of Atropa species varies, with most being relatively secure but some facing significant threats due to their limited distributions, habitat specificity, and historical overcollection for medicinal purposes. Atropa bella-donna, the most widespread species, is generally not considered threatened globally. It is ranked as Apparently Secure to Secure (G4G5) in by NatureServe, though some populations in have declined due to agricultural intensification and targeted eradication efforts. Atropa baetica is listed as Endangered (EN) on the European Red List of Medicinal Plants, under criteria B2ab(iii,iv); D, due to its restricted range in central and southern and northern , with threats including from and , as well as illegal collection. It is protected as a priority under Annex II of the EU Habitats Directive and the Bern Convention. Atropa acuminata, native to the (eastern to and ), is regarded as critically endangered in regional assessments owing to for its alkaloids, habitat , and slow reproductive rates. Conservation efforts include protocols to support ex situ preservation. For Atropa pallidiflora (northern to the ) and Atropa komarovii (northeastern to southern ), formal global conservation assessments are lacking as of 2025, but both are endemic to specific habitats that are under pressure from . Limited populations suggest potential vulnerability, warranting further research and monitoring.

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