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Datura stramonium L.

Protologue Sp. pl. 1: 179 (1753).

Family

Chromosome number 2n = 24

Vernacular names Thorn apple, green thorn apple, Jimson , Jamestown weed, devil’s apple, devil’s trumpet, stramonium (En). Pomme épineuse, stramoine, , feuille du diable, herbe du diable (Fr). Figueira do inferno, pomo espinhoso, erva dos bruxos, palha verde, estramonio (Po). Muranha (Sw).

Origin and geographic distribution is native to the Americas and has been introduced in many tropical, subtropical and even temperate regions. It is a naturalized weed in many African countries, but is probably seriously under-reported.

Uses Datura stramonium and L. have largely similar medicinal uses throughout the world. The most widely known use of Datura stramonium and of other Datura species is for relieving , cough, tuberculosis and bronchitis by smoking the dried , or . ‘Asthma ’ have been shown to be very effective in some cases, but in other cases they had little or no effect. Cigarettes made with the leaves are also used to treat Parkinson’s disease. A decoction or infusion of leaves is given as a sedative to mental and schizophrenic patients. The leaves are applied as a dressing to cure rheumatic pain, swellings, wounds, gout, burns, ingrown toe-nails, fungal infections, tumours and ulcers. Dried pulverized leaves are dusted on wounds or applied after mixing the powder with fat or Vaseline. In DR Congo pounded fresh and fresh leaves are soaked in water and the liquid is given in enema as an abortifacient. In Zimbabwe a hot poultice of leaves and roots is applied to goitre. A infusion is drunk to treat venereal diseases; to cure ulcers the skin is washed with an infusion of roots and leaves. In Burundi leaf ash is eaten as a cure for whooping cough. In Rwanda a leaf infusion is taken as an antispasmodic and to reduce stomach acidity. In Kenya dried and ground leaves and are eaten mixed with fat to treat ringworm. Headache is relieved by rubbing the scalp with leaves or leaf sap. Hair loss is countered by applying sap or leaf pulp and these also serve to remedy dandruff. In Ethiopia pieces of young fruit are sucked against tonsillitis and sore throat and applied to abscesses and swollen glands. In Kenya and Lesotho the fruit is heated in hot ash and after cooling juice is squeezed and used as ear drops to treat earache. In Zimbabwe an infusion of fruit ash is drunk to treat stomach- ache. In Ethiopia the smoke of burning seeds is inhaled to relieve toothache, while in Kenya fresh green fruit is applied for this purpose. In Namibia a leaf extract is administered to cows to ensure a rapid expulsion of the afterbirth and pulped roots are mixed with water and given to cattle to cure lung diseases. The dried leaves and seeds of Datura stramonium are included in the pharmacopoeias of many Western countries as an antispasmodic and for treatment of asthma, whooping cough and Parkinson’s disease. The use of Datura stramonium varies between cultures. In Central and South America hallucinogenic uses are common among native tribes. In Africa, before they enter fighting contests, young men of the Fulfulde people of the border area of Niger and Nigeria are served drinks containing Datura seeds. This increases their courage and pain tolerance. The leaves are most commonly used as a narcotic, either smoked or boiled and eaten; seeds are similarly used. Roots, seeds or leaves are added to alcoholic drinks to increase the intoxicating effect. Side effects include dry mouth and throat, eye pain, blurred vision, restlessness, dizziness, arrythmia, flushing and faintness. An overdose will cause headache, nausea, vomiting and affect the central nervous system causing disorientation, , , inappropriate affect, short-term memory loss and coma. The seeds are also used for criminal purposes. Hospital admissions and fatalities, most often of adolescents, are not uncommon. It is for this reason that several countries including France removed datura cigarettes from the Pharmacopoeia in 1992. Reports on the use of the as an insecticide vary from good control of aphids in crops in Namibia to no effects in Australia. In East Africa the leaves yield a green dye that is used to dye cloth; in Lesotho the twigs yield a blue-green dye that is used for house decoration. In Ethiopia the plant has been used to tattoo the gums, partly as a treatment of gingivitis or dental decay. The stems are used as firewood. In Kenya the oil is used as massage oil.

Production and international trade In tropical Africa Datura stramonium is mainly used locally, but it is important for the international pharmaceutical industry. For example, in France 20–30 t of leaves were used around 1990 to produce anti-asthma and antispasm and medicines against Parkinson’s disease. There are many brand names for and on the world market.

Properties The concentration of total in the leaves of Datura stramonium is 0.2– 0.5%, hyoscyamine being the major compound and (= hyoscine), apoatropine, tropine, belladonnine and hyoscyamine N-oxide minor compounds; more than 70 alkaloids have been identified in the various parts of the plant. Biosynthetically, the main compounds all belong to the alkaloids and are derived from the amino acid ornithine. Hyoscyamine, atropine and scopolamine are , specifically antimuscarinics. They act by competitively and reversibly inhibiting the neuro- transmitter from binding to its muscarinic receptors, and this antagonism leads to sympathomimetic-like effects in the organs. They increase the heart rate, induce relaxation and motor inhibition in smooth muscles, decrease secretions, and induce dilation of the pupils of the eyes. Although hyoscyamine has a stronger activity than atropine or scopolamine, atropine is more commonly prepared and used. Although at low doses their action tends to be and sedative, at high doses they cause substantial excitation: agitation, disorientation, exaggerated reflexes, hallucinations, , mental and insomnia. Hyoscyamine is used to provide symptomatic relief of various gastrointestinal disorders including spasms, peptic ulcers, irritable bowel syndrome, pancreatitis, colic and of the bladder. It has also been used to relieve some heart problems, to reduce excess saliva production and control some of the symptoms of Parkinson’s disease. The Datura stramonium powder listed in the Dutch Pharmacopoeia (8th edition) is titrated to contain 0.23–0.27% total alkaloids. It is an ingredient of antitussive syrups, but is mostly used in the form of cigarettes to relieve respiratory difficulties, together with other . Tropane alkaloids can be biosynthesized in cell suspension cultures of Datura stramonium in shake flasks and bioreactors. Calluses have been induced from leaves, stems and roots and cultured on Gamborg’s B5 or Murashige and Skoog medium supplemented with growth regulators. The highest content was produced in leaf calluses grown on a medium with low concentrations of growth regulators (0.1 mg/ml of benzyladenine and 2,4-D), and in cultures grown in the dark. Total alkaloid production in a cell culture supplemented with phenylalanine and ornithine was 5 times higher than in the control culture, and higher ratios of tropine to tropic acid also stimulated alkaloid production. A hyoscyamine production of up to 7.5 mg/l daily was recorded in root cultures on Gamborg’s B5 medium containing 5% sucrose at 20–25°C. Methanol leaf extracts showed slight antibacterial activity against gram-positive bacteria in a dose dependent manner but no activity was found against Escherichia coli and Pseudomonas aeruginosa. The seed contains about 17% of a pale yellow oil.

Adulterations and substitutes Tropane alkaloids similar to those found in Datura are known from numerous Solanaceae (e.g. hyoscyamine and scopolamine in L. and Hyoscyamus muticus L.). Scopolamine is found in high quantity in in Duboisia spp., which are used for industrial production in Australia.

Description Annual or short-lived perennial erect herb up to 2 m tall, often much-branched; stem sparsely hairy to glabrous. Leaves alternate, simple, minutely hairy; stipules absent; petiole up to 9.5 cm long; blade ovate to rhombic-ovate or elliptical, 3–20 cm × 1–15 cm, base cuneate, rounded, truncate or cordate, apex acute to acuminate or obtuse, margins sharply toothed with irregular teeth or almost entire, pinnately veined. Flowers axillary, solitary, rarely paired, bisexual, regular, 5-merous; pedicel 5–15 mm long, up to 30 mm long in fruit; calyx tubular, 2.5–5 cm long, lobes unequal, 0.5–1 cm long; corolla trumpet-shaped to tubular, 6–10 cm long, white or faintly tinged purple, sometimes violet or purplish in the tube; stamens inserted above the middle of the corolla tube, included, filaments short and thick, anthers yellow; ovary superior, 2(–4)-celled, style slender, 3.5–7 cm long, stigma large, 2-lobed. Fruit an upright, almost globose to ovoid up to 5 cm × 4.5 cm, yellowish to brown, spines few to many, slender, stiff, up to 16 mm long, many-seeded. Seeds almost D-shaped, flattened, 3.5–4.5 mm × 2.5– 3.5 mm × c. 1 mm, dark brown to black. Seedling with epigeal germination; cotyledons thin, leafy.

Other botanical information Datura comprises about 10 species, which all originated in the ; most species have been introduced throughout the world. Datura stramonium belongs to section Stramonium. L. (longspine thornapple, fierce thornapple) belongs to the same section, and is recorded with certainty only in Cape Verde, Namibia, Botswana and Zimbabwe. Compared to Datura stramonium, it has wider leaves and larger with fewer but stouter spines. The medicinal uses are similar to those for the other Datura species. The main alkaloid in its leaves is scopolamine and not hyoscyamine as in Datura stramonium, although both species produce hyoscyamine in the roots. In Datura ferox hyoscyamine is transformed into scopolamine in the above-ground parts. is considered here as a separate , although it is often treated as a section of Datura (section Brugmansia). It mainly differs in its habit (a woody, comparatively long-lived or small ), its mode of growth (reproducing vegetatively by root suckers), its pendulous or inclined flowers open throughout anthesis for 4–6 days with spathe-like, not circumscissile calyx and long pedicel, and its fruit being a usually indehiscent, unarmed berry. Chemically, Datura and Brugmansia are similar, and consequently they have similar medicinal applications. However, the primary use of the Brugmansia species is ornamental.

Growth and development Only the basal part of the stem remains vegetative; flowering occurs on the branched part of the plant and branches do not resume vegetative growth after flowering and fruiting. The flowers are closed during the day and open in the evening, and are reported to be pollinated by hawk and to be largely self- fertile. The hyoscyamine/scopolamine ratio in Datura stramonium is influenced by the developmental stage reached by the . In younger plants scopolamine is the main alkaloid, whereas hyoscyamine mostly becomes the dominant alkaloid when development has started. The alkaloids are produced in the roots and transferred to the leaves, flowers and finally the fruits.

Ecology Datura stramonium occurs in open locations such as grassland, roadsides, waste places, scrub vegetation and open forest. It tolerates various soil types but prefers clayey or loamy soils. Datura stramonium is frost sensitive. In the United States and Australia Datura stramonium is considered a serious weed in crops; elsewhere it is considered a weed in waste land. Control is difficult as Datura stramonium is resistant to most commonly used herbicides. Contamination of wheat, rye, buckwheat and linseed with seeds of Datura stramonium resulting in poisoning have been reported.

Propagation and planting Datura stramonium is generally cultivated from seed sown either directly in the field or in a nursery bed. Soaking seed overnight improves germination. Per ha, 7–8 kg of seed is needed. Seed starts germinating after about 2 weeks, and germination is complete after one month. If the seed is sown in a nursery, seedlings are transplanted when 8–12 cm tall. Normal spacing is 70–100 cm, but in a spacing of 3 m is common practice.

Management Experiments in Burundi with the cultivation of Datura stramonium showed that the application of chemical and manure resulted in an increased production of total alkaloids. As a result of experiments in Burkina Faso, deflowering was recommended to increase the total alkaloid content in the leaves.

Diseases and pests Many pests and diseases affecting solanaceous crops also affect Datura stramonium.

Harvesting In experiments in Burundi it was demonstrated that the best time for harvesting leaves of Datura stramonium was 8 weeks after sowing, because alkaloid content was then highest. On the basis of experiments in Burkina Faso, it was recommended to harvest the leaves in the early morning or late afternoon.

Yield The highest yields of scopolamine and hyoscyamine from Datura stramonium in Algeria were 7.5 kg/ha and 21 kg/ha, respectively. In India 1–1.5 t/ha of dry leaves and 500–600 kg/ha of seeds have been obtained.

Handling after harvest The tender branches and leaves are dried in the shade. Fruits are dried in the sun until they dehisce. The seeds are separated by threshing and after further drying packed for transport.

Genetic resources Datura stramonium has a wide geographical distribution, prefers anthropogenic habitats and is therefore not liable to genetic erosion.

Breeding No attempts have been made to improve Datura stramonium for yield of alkaloids but the genetics of Datura spp. have been extensively studied. Interspecific crosses with Datura ferox yield F1-plants that have the capability to transform hyoscyamine into scopolamine; this characteristic is dominant and monofactorial.

Prospects Although the tropane alkaloids scopolamine, hyoscyamine and atropine can be produced synthetically, it is more economical to extract them from plants such as Datura spp. However, Atropa, Duboisia and Hyoscyamus species are the major sources of raw materials. On the world market of pure tropane alkaloids, it will be difficult for African producers to compete with producers in China, India and Australia.

Major references • Berkov, S., Zayed, R. & Doncheva, T., 2006. Alkaloid patterns in some varieties of Datura stramonium. Fitoterapia 77(3): 179–182. • Burkill, H.M., 2000. The useful plants of West Tropical Africa. 2nd Edition. Volume 5, Families S–Z, Addenda. Royal Botanic Gardens, Kew, Richmond, United Kingdom. 686 pp. • Dethier, M., Cordier, Y. & Demeyer, K., 1993. Cultivation of Datura species for scopolamine and hyoscyamine production in Burundi. Acta Horticulturae 331: 39–48. • Gonçalves, A.E., 2005. Solanaceae. In: Pope, G.V., Polhill, R.M. & Martins, E.S. (Editors). Flora Zambesiaca. Volume 8, part 4. Royal Botanic Gardens, Kew, Richmond, United Kingdom. 124 pp. • Neuwinger, H.D., 2000. African traditional : a dictionary of plant use and applications. Medpharm Scientific, Stuttgart, Germany. 589 pp. • Nkurunziza, J.P., undated. Validating traditional knowledge: Rwanda. In: Sharing innovative 10: Examples of the development of pharmaceutical products from . UNDP & TWNSO, Trieste, Italy. pp. 127–134. • Philipov, S., Berkov, S. & Doncheva, T.S., 2007. GC-MS survey of Datura stramonium alkaloids. Comptes Rendus de l’Académie Bulgare des Sciences 60(3): 239–250. • Sri Hartati, Imastini Dinuriah & Blomqvist, M.M., 1999. Datura L. In: de Padua, L.S., Bunyapraphatsara, N. & Lemmens, R.H.M.J. (Editors). Plant Resources of South-East Asia No 12(1). Medicinal and poisonous plants 1. Backhuys Publishers, Leiden, Netherlands. pp. 229–234.

Other references • Abebe, D. & Hagos, E., 1991. Plants as a primary source of drugs in the traditional health practices of Ethiopia. In: Engels, J.M.M., Hawkes, J.G. & Worede, M. (Editors). Plant genetic resources of Ethiopia. Cambridge University Press, Cambridge, United Kingdom. pp. 101–113. • Al-Shaikh, A.M. & Sablay, Z.M., 2005. Hallucinogenic plant poisoning in children. Saudi Medical Journal 26(1): 118–121. • Blomqvist, M.M., 1997. and uses of medicinally important species in the genera Datura L. and Solanum L. (Solanaceae) in South East Asia. Unpublished MSc. thesis, Department of Plant Taxonomy, Wageningen Agricultural University, the Netherlands. 132 pp. • Braun, M., Burgstaller, H., Hamdoun, A.M. & Walter, H., 1991. Common of Central Sudan. Margraf, Weikersheim, Germany. 329 pp. • Eftekhar, F., Yousefzadi, M. & Tafakori, V., 2005. Antimicrobial activity of and Datura stramonium. Fitoterapia 76: 118–120. • Fortin, D., Lô, M. & Maynart, G., 1990. Plantes médicinales du Sahel. ENDA, Dakar, Senegal & CECI, Montréal, Canada. 280 pp. • Friedman, M., 2004. Analysis of biologically active compounds in potatoes (Solanum tuberosum), tomatoes (Lycopersicon esculentum), and jimson weed ( Datura stramonium) seeds. Journal of Chromatography 1054(1–2): 143–155. • Gedif, T. & Hahn, H.-J., 2003. The use of medicinal plants in self-care in rural central Ethiopia. Journal of Ethnopharmacology 87: 155–161. • Gelfand, M., Mavi, S., Drummond, R.B. & Ndemera, B., 1985. The traditional medical practitioner in Zimbabwe: his principles of practice and pharmacopoeia. Mambo Press, Gweru, Zimbabwe. 411 pp. • Gonçalves, A.E., 2002. Solanaceae. In: Martins, E.S., Diniz, M.A., Paiva, J., Gomes, I. & Gomes, S. (Editors). Flora de Cabo Verde: Plantas vasculares. No 71. Instituto de Investigação Científica Tropical, Lisbon, Portugal & Instituto Nacional de Investigação e Desenvolvimento Agrário, Praia, Cape Verde. 71 pp. • Gurib-Fakim, A., Sewraj, M., Guého, J. & Dulloo, E., 1993. Medical of some weeds of Mauritius and Rodrigues. Journal of Ethnopharmacology 39(3): 177–185. • Hamill, F.A., Apio, S., Mubiru, N.K., Mosango, M., Bukenya-Ziraba, R., Maganyi, O.W. & Soejarto, D.D., 2000. Traditional herbal drugs of southern Uganda, 1. Journal of Ethnopharmacology 70: 281–300. • Hegnauer, R., 1973. Chemotaxonomie der Pflanzen. Band 6. Birkhäuser Verlag, Basel, Switzerland. 882 pp. • Maundu, P., Berger, D., Saitabau, C., Nasieku, J., Kipelian, M., Mathenge, S., Morimoto, Y. & Höft, R., 2001. Ethnobotany of the Loita Maasai. Towards community management of the forest of the Lost Child. Experiences from the Loita Ethnobotany Project. UNESCO People and Plants Working Paper 8, Paris, France. 34 pp. • Nwosu, M.O., 1999. Herbs for mental disorders. Fitoterapia 70: 58–63. • Reisman-Berman, O., Kigel, J. & Rubin, B., 1988. Factors involved in the germination of Datura ferox and D. stramonium. Phytoparasitica 16(4): 371–372. • Roddick, J.G., 1991., 1991. The importance of the Solanaceae in medicine and therapy. In: Hawkes, J.G., Lester, R.N., Nee, M. & Estrada, R.N. (Editors). Solanaceae 3. Taxonomy, chemistry, evolution. Royal Botanic Gardens, Kew, Richmond, United Kingdom. pp. 7–23.

Sources of illustration • DeWolf, G.P., 1956. Notes on cultivated Solanaceae 2: Datura. Baileya 4(1): 13–23. • Gonçalves, A.E., 2005. Solanaceae. In: Pope, G.V., Polhill, R.M. & Martins, E.S. (Editors). Flora Zambesiaca. Volume 8, part 4. Royal Botanic Gardens, Kew, Richmond, United Kingdom. 124 pp.

Author(s) • F.S. Mairura Kenya Tropical Soil Biology and Fertility Institute of CIAT, P.O. Box 30677, Nairobi, Kenya

• M.P. Setshogo Department of Biological Sciences, University of Botswana Herbarium, Private Bag UB00704, Gaborone, Botswana

Editors • G.H. Schmelzer PROTA Network Office , Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands

• A. Gurib-Fakim Faculty of Science, University of Mauritius, Réduit, Mauritius Associate editors • C.H. Bosch PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands

• M.S.J. Simmonds Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, United Kingdom

• R. Arroo Leicester School of Pharmacy, Natural Products Research, De Montfort University, The Gateway, Leicester LE1 9BH, United Kingdom • A. de Ruijter PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands General editors • R.H.M.J. Lemmens PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands

• L.P.A. Oyen PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands Photo editor • A. de Ruijter PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands

naturalized

1, flowering branch; 2, fruit. Redrawn and adapted by Iskak Syamsudin

plant habit obtained from Plants of Hawaii

Plant habit obtained from Plants of Hawaii

flowering branch obtained from malezasdemexico

fruit obtained from P.J. Alexander

dry fruit obtained from Plants of Hawaii

opened fruit obtained from Plants of Hawaii

seeds obtained from S. Hurst