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Gloriosa superba L.

Protologue
Sp. pl. 1: 305 (1753)
Family
Colchicaceae
Chromosome number
2n = 14, 22, 33, 44, 66, 77, 84, 88, 90
Synonyms
Gloriosa simplex L. (1767), Gloriosa virescens Lindl. (1825), Gloriosa abyssinica A.Rich. (1850), Gloriosa carsonii Baker. (1895), Gloriosa minor Rendle (1896), Gloriosa baudii (N.Terracc.) Chiov. (1916).
Vernacular names
Glory lily, flame lily, climbing lily, creeping lily (En). Lis de Malabar, lis grimpant, lis glorieux (Fr). Garras de tigre, aranha de emposse (Po). Mkalamu, kimanja nouchawi (Sw).
Origin and geographic distribution
Gloriosa superba occurs naturally in Africa, in India, and southeastern Asia, and is nowadays distributed widely throughout the tropics, and worldwide as a pot plant. In Africa, its distribution is from Senegal east to Ethiopia and Somalia, and south to South Africa.
Uses
In tropical Africa the different parts of Gloriosa superba have a wide variety of uses, especially in traditional medicine. In Côte d’Ivoire a leaf decoction applied as a liniment eases cough and general pain, and leaf juice is instilled into the nose in case of fainting. In Côte d’Ivoire and Burkina Faso leaves are administered in enema as a decongestant. In Congo crushed leaves are applied to the chest to treat asthma. In Burundi a leaf decoction is recommended for treating dropsy of the scrotum, while the leaf pulp serves against rheumatism. The Ulanga people of Tanzania burn the herb and apply ash on wounds to promote healing. They also drink the plant juice as an antimalarial.
At low doses, the tuber has numerous medicinal applications. It is used traditionally for the treatment of bruises, colic, chronic ulcers, haemorrhoids and cancer, and is also employed as a tonic and purgative. It is put into poultices to relieve neuralgia, and used in topical applications to treat arthritic conditions, swellings of the joints, sprains and dislocations. The tuber is claimed to have antidotal properties to snakebites. In Sudan tuber sap is an ingredient of a drink that induces sleep. The Marakwet people of Kenya take a tuber decoction against abdominal disorders and to induce abortion. Macerated tuber is also taken against smallpox, leprosy, eczema, itch, and ringworm. In DR Congo the rasped and washed tuber is used externally to treat venereal diseases and stomach-ache.
The anthelmintic properties of the tuber, fruits and leaves are widely known and they are used to treat infections of Guinea worms, schistosomes (causing bilharzia), roundworm, tapeworm, liver fluke and filaria. A rectal injection of the juice from mashed leaves is applied to cure female sterility among Pygmy groups. A paste made from the tuber is applied externally to facilitate parturition. The Ulanga people of Tanzania use tuber juice for ear drops to treat earache, while the Shona people of Zimbabwe drop tuber juice on painful teeth. In Zambia the tuber is part of a preparation for impotence, and is used as an abortifacient. Soup made from leaf or tuber sap is given to women suffering from sterility, delayed puberty, delayed childbirth and menstrual problems. Leaf juice, unripe fruits mixed with butter, and tuber macerate are frequently used to kill head lice.
In northern Nigeria the tuber is added to arrow poison based on Strophanthus sp. In coastal Kenya and Tanzania, powdered tuber is commonly used as a suicidal agent and to commit homicide, because of its high toxicity. The species is also widely believed to have magical properties.
Several cultivars of Gloriosa superba are cultivated in the tropics and under greenhouse conditions in temperate regions, the commonest being ‘Rothschildiana’. It is grown both as a cut flower and as a pot plant.
Production and international trade
Gloriosa superba is exported by India and Sri Lanka to pharmaceutical industries, and more recently also by a few African companies based in Nigeria, Cameroon and Zimbabwe. Quantities involved are not known.
Properties
The medicinal importance of Gloriosa superba is due to the presence of alkaloids in all parts of the plant, mainly colchicine (superbine), an amino alkaloid derived from the amino acids phenylalanine and tyrosine. The presence of colchicine-type alkaloids with a tropolone ring is characteristic for most genera in Colchicaceae. The seeds are the best source of colchicine, as their content is 2–5 times higher than in the tubers. A report from Rwanda claims that the highest colchicine content is found in young leaves.
Several colchicine-related alkaloids have been isolated from tubers and seeds. They are mostly demethyl substitutes and include cornigerine, which is a potent antimitotic, and colchicoside used as a muscle relaxant. A plant can contain up to 0.9% colchicine and 0.8% colchicoside.
In medicine, colchicine is used in the treatment of gout. In spite of its serious side-effects it is still commonly used for acute gout. It reduces the inflammatory reaction to urate crystals deposited in the joints. Its efficacy might be due to decreased leukocyte mobility. The substance is not an analgesic, and has no effect on blood concentration, nor on renal excretion of uric acid. Diarrhoea, nausea, vomiting and abdominal pains are the first signs of poisoning and occur at doses equal to or lower than those needed to treat gout. The diarrhoea may become severe and haemorrhagic. A burning sensation in the throat, stomach and skin may also be an early sign of intoxication. Nibbling on the tubers causes numbness of the lips and tongue and loss of body hair. Severe reactions include extensive vascular damage and acute renal toxicity with oliguria and haematuria. The patients may develop convulsions, delirium, muscle weakness, neuropathy and ascending paralysis of the central nervous system. In patients who have taken an overdose of Gloriosa superba tubers, death occurs as a result of respiratory depression and cardiovascular collapse within a few days.
Colchicine is a powerful antimitotic agent that blocks or suppresses cell division by inhibiting mitosis, the division of a cell’s nucleus. It is used in plant breeding to induce polyploidy, as it allows chromosome division but inhibits formation of a mitotic spindle figure, which guides the separation of the two sets of haploid chromosomes. As a result no sister cells are formed. Once the treatment has stopped, however, the normal spindle figure forms again. Colchicine also inhibits the division of animal cells, but it is too toxic to be used to arrest tumour growth. A biosynthetic precursor of colchicine, demecolcine, has a wider margin of safety and is used to treat myelogenic leukaemia and malignant lymphoma. Extracts of the shoots and of the tubers of Gloriosa superba show strong nematicidal activity, which can be largely attributed to colchicine. The chemical constituents of the tuber are known to be very poisonous to fish. Severe damage is done by colchicine-containing plants to livestock in different parts of Africa.
In vitro production of colchicine is feasible, although the levels are in general 10–25 times lower than those found in plants grown in vivo.
Adulterations and substitutes
The corms of Colchicum autumnale L. and Iphigenia spp. (also Colchicaceae) are traditional sources of colchicine. An increase in demand for colchicine stimulated the search for an alternative source, leading to Gloriosa superba. Chemical synthesis of colchicine is possible but complicated. Synthesis is expected to remain an important target of the chemical industry. In-vitro production of colchicine is also possible but gives low yields.
Description
Climbing, sometimes erect herb up to 4 m long; stem annual, glabrous and sparsely branched; tuber perennial, horizontal, abruptly bent in a V or L shape, roots fibrous. Leaves in whorls of 3–4, opposite or alternate, simple, sessile; blade ovate to lanceolate, 6–15(–20) cm × 1.5–4 cm, base obtuse, apex of upper leaves with or without 1–2 cm long tendril, parallel-veined. Flowers axillary, solitary, bisexual, regular, 6-merous, 4.5–7 cm in diameter, showy, pendulous; pedicel 4–20 cm long; perianth segments free, lanceolate or oblanceolate, 5–7(–9) cm × 1(–2) cm, often with undulate margins, strongly reflexed when mature, persistent, usually yellow and red, less often yellow, red or white; stamens with filaments 2–5 cm long, spreading, anthers 7–10 mm long, opening by longitudinal slits; ovary superior, 3-celled, carpels coherent only by their inner margins, style filiform, 2–4(–5.5) cm long, bent at a right angle basally. Fruit a loculicidal, oblong capsule 4–6 cm × 1–2 cm, containing up to 20(–40) seeds. Seeds ovoid, 4–5 mm in diameter, surrounded by a fleshy, red sarcotesta.
Other botanical information
The taxonomy of Gloriosa is confused, and up to 27 species have been recognized. Gloriosa superba is considered here a single highly variable species. In Zimbabwe morphologically uniform populations occur which have variable polyploidy levels, but the cytological differentiation does not reflect any precise geographical trend. The widely cultivated cultivar ‘Rothschildiana’ is hexaploid.
Growth and development
Gloriosa superba is a typical geophyte whose aerial stems die down in the dry season and the tuber is dormant during this period, only sprouting with the rains. Two or more tubers develop during each growing season, while the previous season’s tuber starts to shrivel. The tuber contains mainly starch, which increases gradually until the tuber is full grown. Colchicine content in tubers increases simultaneously. Plants propagated from seeds take 3–4 years to bloom. Plants produced from tubers develop (1–)3–6 stems, which start flowering after 5–8 weeks and continue flowering for about another 7 weeks, after which the stems die. Development from visible flower bud to bloom takes about 2 weeks and anthesis occurs 1 day later with the stigma being receptive for 4 days; anther dehiscence is one day after anthesis. The same branch flowers at 3-day intervals. Terminal flowers do not usually set fruit, but if they do, only a few seeds are produced. Pollination is probably by butterflies and sunbirds. Fruits are mature 6–10 weeks after pollination. The red sarcotesta suggests seed dispersal by animals. The dimensions and branching pattern of the plants are strongly correlated with tuber weight.
Ecology
Gloriosa superba prefers a climate with a pronounced rainy season, avoiding per-humid tropical areas. It is most common in forest-savanna boundaries. It is locally common in thickets, hedges, open forest, grassland and bushland, where it can be seen scrambling through shrubs, and is also found in abandoned cultivated areas. It occurs from sea-level up to 2500 m altitude. Gloriosa superba grows best in well-drained, acid to neutral soil rich in organic matter.
Propagation and planting
Gloriosa superba is propagated mainly during the rainy season, by bulblets, division of the tubers or from seed. V- or L-shaped tubers should be divided every third year. The tuber is delicate, and should be teased apart gently just before new growth begins, when the buds are easiest to spot. Each tuber part must contain several axillary buds that ensure the formation of adventitious stems and roots. Vegetative propagation by tubers is common practice but slow as the maximum number of daughter tubers produced per plant per year is two. Separating 2-lobed tubers produces a higher percentage of flowering plants than leaving the tubers undivided (97% versus 63%). Sprouting of the tubers is irregular and reaches about 60% in 30 days. Tuber dormancy can be overcome by soaking in continuously aerated water. Small tubers have been found to have a higher success rate than bigger ones. Tubers of 50–60 g are planted horizontally and 30–45 cm apart in well-tilled soil at a depth of 6–8 cm in furrows 45–60 cm apart. A closer spacing gives a higher percentage of cross-pollination resulting in improved fruit set. The best planting medium is a 1:1:2 mixture of soil, sand and compost.
Growing Gloriosa superba from seed requires more time. The seeds have to be soaked overnight in warm water, and then planted in a well-drained medium. Germination is erratic and may take from 3 weeks to 3 months. Seedlings grow rapidly and mostly produce tubers by their second year; flowering starts in the fourth year. Chemical scarification (e.g. with 1% hypochlorite) or removal of the sarcotesta reduces seed dormancy from 6–9 months to about 4 months, and accelerates germination to 11–15 days. Germination rates as high as 97% have been reached for seeds incubated at 20–25°C for a period of 31 days. Higher temperatures have adverse effects.
As seed germination is poor and vegetative propagation is slow, methods of rapid micropropagation have been developed, using explants preferably from tubers and an auxin-enriched growth medium. Explants from meristematic parts of the tuber yield the highest number of plantlets. A growth medium containing thiamine hydrochloride and NaCl yielded large numbers of mature microtubers in 3–4 months. Periodic darkness accelerates this process.
Management
Gloriosa superba prefers light to medium shade. In general, the plants are grown organically. Before sowing, 15–20 t of farmyard manure is applied. It is also possible to apply a dose of 40 kg N, 50 kg P2O5 and 75 kg K2O per ha at planting with a top dressing of 80 kg N per ha, 8 weeks after planting. The top dressing should coincide with staking of the growing stems. Irrigation is needed for dry spells during the initial stages of growth. Irrigation applied after flowering may cause the tubers to rot. Poor fruit set is a problem in plantings in southern India. This may be attributed to inadequate pollination, which can be overcome by artificial pollination. Although considerable information is available on the cultivation of Gloriosa superba under greenhouse conditions, the techniques used are not applicable to field conditions in tropical regions.
Different pollination methods were studied including natural pollination, controlled selfing and cross-pollination. Although flower colour and shape seem to favour cross-pollination, self-pollination provides better results. Controlled selfing between flowers on the same plant (idiogamy) gives significantly higher seed yield (9.2 g/plant), compared to naturally pollinated ones (4.3 g/plant).
Diseases and pests
Leaf blight (Curvularia lunata) and tuber rot (Sclerotium spp.) are important fungal diseases of Gloriosa superba under per-humid conditions. Caterpillars of the moths Polytela gloriosa and Chrysodeixis chalcites attack foliage and flower buds. Under greenhouse conditions, lice and thrips can be a problem.
Harvesting
Mature fruits of Gloriosa superba are hand picked, and the tubers are dug out manually.
Yield
In South Africa the seed production of ‘wild-type’ plants is positively correlated with height of the plant, and is on average 258 seeds per plant for plants 60–65 cm tall compared with about 30 seeds per plant for plants 30–40 cm tall. In Tamil Nadu, India, small-scale plantings, raised from tubers, yield on average 250–300 kg seed/ha from the second year onwards.
Handling after harvest
Seeds and tubers are traded in various forms: fresh, dry, powdered, or in oil. In South-East Asia, after harvesting, mature fruits of Gloriosa superba are left in the shade to dry for 7–10 days. The fruits are then split open and the seeds removed, dried for a week in the shade and subsequently sun dried for another week.
Genetic resources
Gloriosa superba has a wide natural distribution, and many selections are cultivated. Local depletion of the resource does occur, in India particularly, where the species has become endangered due to overcollection of the tubers. Although in its natural habitat, seed set and germination is poor. Gloriosa superba is not threatened and the diversity still offers opportunities for further selection either for chemical constituents or as an ornamental. Investment analysis shows that Gloriosa superba cultivation is profitable under both irrigated and rainfed conditions. There are no known germplasm collections of Gloriosa superba.
Breeding
There are several cultivars of Gloriosa superba for ornamental purposes, with ‘Rothschildiana’ being the most frequently encountered; ‘Lutea’ is a yellow flowered form, while ‘Citrina’ is yellow with dark red markings. There is also a dwarf cultivar called ‘Nana’.
Prospects
Gloriosa superba is widely used as a medicinal plant in Africa, despite the fact that the whole plant is very poisonous. It is therefore recommended that protocols for safe use of the different plant parts be developed. Gloriosa superba is commonly grown as a garden ornamental in the tropics and also shows interesting potential as a source of colchicine. Thus, in Africa there might be good opportunities for planting the crop commercially for ornamental purposes, or for the production of colchicine.
Major references
• Bunyapraphatsara, N. & van Valkenburg, J.L.C.H., 1999. Gloriosa superba 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. 289–292.
• Burkill, H.M., 1995. The useful plants of West Tropical Africa. 2nd Edition. Volume 3, Families J–L. Royal Botanic Gardens, Kew, Richmond, United Kingdom. 857 pp.
• Duke, J.A., 1992. Handbook of phytochemical constituents of GRAS herbs and other economic plants. CRC Press, Boca Raton, Florida, United States. 654 pp.
• Finnie, J.F. & Van Staden, J., 1994. Gloriosa superba L. (Flame Lily): Micropropagation and in vitro production of colchicine. In: Bajaj, Y.P.S. (Editor): Biotechnology in agriculture and forestry. Volume 26. Medicinal and Aromatic Plants VI. Springer Verlag. Berlin, Heidelberg, New York, London, Paris, Tokyo. pp. 147–166.
• Iwu, M.M., 1993. Handbook of African medicinal plants. CRC Press, Boca Raton, Florida, United States. 464 pp.
• Le Roux, L.G. & Robbertse, P.J., 1994. Tuber ontogeny, morphology and vegetative reproduction of Gloriosa superba L. South African Journal of Botany 60: 321–324.
• Le Roux, L.G. & Robbertse, P.J., 1997. Aspects relating to seed production in Gloriosa superba L. South African Journal of Botany 63(4): 191–197.
• Neuwinger, H.D., 1996. African ethnobotany: poisons and drugs. Chapman & Hall, London, United Kingdom. 941 pp.
• Raina, R. & Gupta, L.M., 1999. Increasing seed yield in glory lily (Gloriosa superba) - experimental approaches. Acta Horticulturae 502: 175–179.
• Sivakumar, G. & Krishnamurthy, K.V., 2000. Micropropagation of Gloriosa superba L., an endangered species of Asia and Africa. Current Science 78(1): 647–659.
Other references
• Chumsri, P., Sawangsuksakul, P., Sornchai, S. & Tempeam, A., 2000. Bulbil formations from plantlets of Gloriosa superba L. Mahidol University Annual Research Abstracts 28: 186.
• Cortnummé, J., Wehrenfennig, M. & Horn, W., 1997. Temperaturbehandlung im Lager von Gloriosa superba (Liliaceae). Poster, Jahrestagung der Deutschen Gartenbauwissenschaftlichen Gesellschaft 1996, Hannover. BDGL-Schriftenreihe 14. p. 138.
• Engprasert, S., 1995. Isolation, structure elucidation, assay and cytotoxic property of tropolone alkaloids from tubers of Gloriosa superba Linn. MSc Thesis, Faculty of Graduate Studies, Mahidol University, Bangkok, Thailand.
• Farooqi, A.A., Kumaraswamy, B.K., Bojappa, K.N., Pusalkar, V.R. & Gupta, R., 1993. Plantations of the clinically important Gloriosa superba. Indian Horticulture 37(4): 26–29.
• Fernando, R. & Widyaratna, D., 1997. Gloriosa superba L. [Internet] International Programme on Chemical Safety (IPCS), Chemical Safety Information from Intergovernmental Organizations, INCHEM. http://www.inchem.org/documents/pims/plant/pim245.htm. Accessed 27 July 2004.
• Field, D.V., 1972. The genus Gloriosa. Lilies 1973: 93-95.
• Khumwanich, K., 1999. Effect of crude extract from climbing lily Gloriosa superba Linn. seeds on polyploid induction of watermelon Citrullus lanatus Mats & Nakai in vitro. MSc Thesis, Chulalongkorn University, Bangkok, Thailand. 90 pp.
• Maroyi, A., 1999. Taxonomic studies on the family Colchicaceae in Zimbabwe with emphasis on variation in the genus Gloriosa. M.Ph. Thesis, Department of Biological Sciences, Faculty of Science, University of Zimbabwe, Harare, Zimbabwe. 117 pp.
• Ntahomvukiye, D., Hakizimana, A., Nkiliza, J. & Van Puyvelde, L., 1984. Dosage de la colchicine dans le Gloriosa simplex L. (Liliacées) du Rwanda (Afrique Centrale). Plantes Médicinales et Phytothérapie 18(1): 24-27.
• Pandey, R. & Haseeb, A., 1988. Studies in the toxicity of extracts of certain medicinal plants to rootknot nematode Meloidogyne incognita (Kifoid & White) Chitwood. Indian Journal of Plant Pathology 6(2): 184–186.
• Saravanan, S. & Buvaneswaran, C., 2003. Gloriosa superba L. cultivation in Tamil Nadu. A socio-economic analysis. Advances in Plant Sciences 16(1): 23–28.
• Sivakumar, G. & Krishnamurthy, K.V., 2002. Gloriosa superba L. - a very useful medicinal plant. In: Singh, V.K., Govil, J.N., Hashmi, S. & Singh, G. (Editors). Recent progress in medicinal plants Volume 7: Ethnomedicine & Pharmacognosy II. Sci-Tech. Publishing LLC, Houston, Texas, USA. pp. 465–481.
• Tarar, J.L. & Vishwakarma, M., 1995. Chromosome morphology of diploid and tetraploid Gloriosa superba Linn. In: Padhye, M.D., Mukherjee, P.K. & Khalatkar, A.S. (Editors). Botany Towards 2000 A.D. (Prof. V.R. Dnyansagar Commemoration Volume). Vedams Book, New Delhi, India.
• van Wyk, B.E., van Heerden, F. & van Oudtshoorn, B., 2002. Poisonous plants of South Africa. Briza Publications, Pretoria, South Africa. 288 pp.
• Watt, J.M. & Breyer-Brandwijk, M.G., 1962. The medicinal and poisonous plants of southern and eastern Africa. 2nd Edition. E. and S. Livingstone, London, United Kingdom. 1457 pp.
Sources of illustration
• Bunyapraphatsara, N. & van Valkenburg, J.L.C.H., 1999. Gloriosa superba 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. 289–292.
Author(s)
E. Dounias
Centre d’Ecologie Fonctionnelle et Evolutive, Center for Evolutionary and Functional Ecology (CEFE-CNRS), 1919, route de Mende, 34293 Montpellier cedex 5, France
Based on PROSEA 12(1): ‘Medicinal and poisonous plants 1’.

Editors
G.H. Schmelzer
PROTA Network Office Europe, 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

Correct citation of this article:
Dounias, E., 2006. Gloriosa superba L. In: Schmelzer, G.H. & Gurib-Fakim, A. (Editors). Prota 11(1): Medicinal plants/Plantes médicinales 1. [CD-Rom]. PROTA, Wageningen, Netherlands.
Distribution Map wild


1, flowering stem; 2, tubers; 3, fruit.
Source: PROSEA



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