Gloriosa superba (PROSEA)

Plant Resources of South-East Asia Introduction

List of species

1, flowering stem; 2, tubers; 3, fruit (Iskak Syamsudin)

Gloriosa superba L.

Protologue: Sp. pl. 1: 305 (1753).

Family: Colchicaceae

Chromosome number: 2n= 22, 66, 88

• Gloriosa simplex auct.

• Flame lily, superb lily (En)
• Indonesia: kembang sungsang (general), pacing tawa (Javanese), katongkat (Sundanese)
• Philippines: climbing lily, Turk's cap
• Cambodia: var sleng dong dang
• Laos: phan ma ha
• Thailand: dong dueng (general), ma khaa kong (northern), waan kaam puu (central)
• Vietnam: cây ngót nghẽo, cây nhú nhoái.

Flame lily is widespread in the Old World tropics, occurring from tropical and southern Africa and Madagascar to India, Burma (Myanmar) and South-East Asia as far as Java and the Lesser Sunda Islands. It is not indigenous in Sumatra and Borneo. It is commonly grown as a garden ornamental in the tropics.

The use of flame lily in folk medicine in South-East Asia is rather limited, probably due to its high toxicity. This may explain the reports of its use as poison in Cambodia and as suicidal agent in Burma (Myanmar). The tubers are applied in traditional Ayurvedic medicine in India. They are used as a tonic, cholagogue, antiperiodic, alterative, anti-arthritis, antileprotic, antihaemorrhoid and purgative. In East Africa the tubers have various applications in folk medicine but their main use exploits the poisonous constituents.

The selection known as G. rothschildiana O'Brien is cultivated in Europe and the United States for the production of cut flowers.

Flame lily seeds are valued as a commercial source of colchicine, an amino alkaloid which is biosynthetically derived from the amino acids phenylalanine and tyrosine. The tubers also contain colchicine, but the content of the seeds is reported to be up to ten times higher, hence the importance attached to good seed set when the plant is grown for colchicine production. A report from Rwanda claims that the highest colchicine content is present in the young leaves.

In medicine, colchicine is used in the treatment of gout and rheumatism. At present it is the drug of choice for acute gout. It reduces the inflammatory reaction to urate crystals deposited in the joints. Its efficacy might be due to several actions, including decreased leucocyte mobility. The substance is not an analgesic, and has no effect on blood concentration, nor renal excretion of uric acid. Because of its highly toxic nature, colchicine should be used under supervision of a physician. Diarrhoea, nausea, vomiting and abdominal pains are often the first signs of poisoning. The diarrhoea may become severe and haemorrhagic, and can thus lead to metabolic acidosis, dehydration, hypotension and shock. A burning sensation in the throat, stomach and skin may also be an early sign of intoxication. 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 G. superba bulbs, death occurs as a result of respiratory depression and cardiovascular collapse.

Colchicine inhibits cell division and is used in plant breeding to produce polyploidy, as it does not prevent chromosome division but inhibits formation of a mitotic spindle figure. Therefore no sister cells are formed. This so-called "C-mitotic" activity of the alkaloid may arise from interaction with the disulphide bonds of the spindle protein, and from inhibition of the conversion of globular proteins to fibrous proteins. Once the treatment has stopped, however, the spindle figure forms again in the normal way. Colchicine also inhibits the division of animal cells, but it is too poisonous to be used to arrest tumour growth. A biosynthetic precursor of colchicine, demecolcine, has a wider margin of safety and is used for the treatment of myelogenic leukaemia and malignant lymphoma.

Extracts of the shoots and of the tubers of the plant 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. Uterine stimulant properties are also reported.

Several other alkaloids have been isolated from tubers and seeds besides colchicine e.g. 1,2-didesmethyl colchicine, 2,3-didesmethyl colchicine, 3-desmethyl colchicine, N-formyl-N-deacetyl colchicine, N-deacetyl-2,3-didesmethyl colchicine, cornigerine, 2-desmethyl colchifoline and colchicoside.

The corms of Colchicum autumnale L. and Iphigenia (both also included in Colchicaceae) are traditional sources of colchicine. An increase in demand for colchicine stimulated the search for an alternative source, leading to G. superba.

• A climbing or, less often, erect, glabrous herb, 3(-6) m long; tuber (corm) perennial, horizontal, roots fibrous; aerial stem annual, moderately branched.
• Leaves alternate, opposite or in whorls of 3(-4), sessile, lanceolate or ovate, entire, base obtuse, apex gradually narrowing in a coiled tendril (except for erect forms).
• Flowers solitary in the axils of leaves, bisexual; perianth segments 6, connate at the base, narrowly elliptical, 5-7(-9) cm × 1(-3) cm, with undulate margins, usually reflexed, yellow, red or 2-coloured; stamens 6, filaments spreading 2.5-5 cm long, anthers 0.7-1 cm long, dorsifixed; ovary superior, sessile, 3-celled, style filiform, bent basally almost at right angle, 3.5-5.5 cm long including short apical branches.
• Fruit a capsule, 4-10 cm × 1.5-2 cm. Seeds globose, about 5 mm in diameter, red or orange red, with a fleshy sarcotesta.

The perennial organ of flame lily should be classified as a hypopodial tuber, mostly consisting of two metaphorous hypopodia with an apical bud enveloped by a prophyll, all covered in the remains of leaf bases. Two or more hypopodial tubers are left in the substrate after each growing season, while the previous season's tuber starts to shrivel. Plants propagated from seed take 3-4 years to bloom. Plants produced from tubers start flowering after 5 weeks, and continue flowering for about another 7 weeks. Development from visible flower bud to bloom takes about 2 weeks and anthesis occurs 1 day later; anther dehiscence takes another day. 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. Cross-pollination generally improves seed production. Pollination is probably by butterflies and sugar birds. Fruit is mature 6-10 weeks after pollination. The dimensions of the plants are strongly correlated with tuber weight, whereas the plants's dimensions determine the size of the next season's tuber.

Gloriosa has often been included in Liliaceae, but is nowadays considered as belonging to Colchinaceae, differing from the bulbous Liliaceae s.s. in having a tuber or corm, i.e. a thickened underground stem part which is stoloniferous in Gloriosa. G. superba is very variable and is sometimes considered to consist of several species. Only one species, G. superba s.s., is indigenous to Malesia. Several selected forms in cultivation are traded under a "specific rank", e.g. G. rothschildiana

In general, flame lily prefers a pronounced monsoon climate, avoiding per-humid tropical areas. It is locally common in brushwood, hedges and open forest up to 300(-600) m altitude. It can be found flowering and fruiting throughout the year, although flowering is most abundant during the rainy season.

Chemical scarification (e.g. with 1% hypochlorite) or removal of the sarcotesta reduces seed dormancy in flame lily from 6-9 months to about 4 months, and accelerates germination from 29-30 days to 11-15 days. Germination rates as high as 97% were reached for seeds incubated at 20-25°C for a period of 31 days. Higher temperatures have adverse effects.

Vegetative propagation by tubers is common practice, but not very suitable for the establishment of large plantings. The maximum number of daughter corms produced per plant is two. Separating the bilobed hypopodial tubers produces a higher percentage of flowering plants than leaving the tubers undivided (about 97% versus about 63%). Tuber dormancy can be overcome by soaking in continuously aerated water. Small tubers have been found to have a higher multiplication rate than bigger ones. Seed yield could be obtained from tubers heavier than 7 g.

Experiments with clonal propagation, using a Murashige and Skoog medium supplemented with 0.1 mg/l indole acetic acid have given promising results. However, the cloned plants did not flower at the first vegetative phase but produced small tubers that could be used as vegetative propagules.

For small-scale plantings, tubers of 50-60 g are planted 30-45 cm apart in well-tilled soil at a depth of 6 cm in furrows 45-60 cm apart. A closer spacing gives a higher percentage of cross-pollination resulting in improved fruit set. The best seeding medium is a 1:1:2 mixture of soil, sand and compost. In Thailand, the optimum fertilizer (13-13-21) rate for optimal seedling growth and tuber weight was found to be about 250 kg/ha.

In vitro production of colchicine is feasible. Levels of colchicine extracted from G. superba callus, malformed roots, and entire plantlets show an increase that can be directly related to the amount of differentiation in culture. To derive levels of colchicine in vitro equal to those found in complete plants, entire plantlet regeneration is necessary.

Although considerable information is available on the cultivation of flame lily under greenhouse conditions, the techniques used are not applicable to field conditions in tropical regions.

At the time of planting a dose of 40 kg N, 50 kg P₂O₅and 75 kg K₂O per ha should be applied with a top dressing of 80 kg N per ha, 8 weeks after planting. The top dressing should coincide with staking of the growing vines. Irrigation is needed at dry weather during the initial stages of growth. Irrigation applied after flowering may cause the tubers to rot.

Poor fruit set resulting in low yield of seed is a problem in plantings in southern India. This may be attributed to inadequate pollination, which can be overcome by artificial hand pollination.

Leaf blight (Curvularia lunata) and tuber rot (Sclerotium spp.) are two important fungal diseases of flame lily under per-humid situations. Lily caterpillar (Polytela gloriosae) and green caterpillar (Pulsia chalcites) attack foliage and flower buds.

Mature fruits of flame lily are hand picked, and the tubers are dug out manually.

In Tamil Nadu, India, small-scale plantings, raised from tubers, yield on average 250-300 kg of seed per ha from the second year onwards. 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.

After harvesting, mature fruits of flame lily 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.

G. superba has a wide natural distribution, and many selections are cultivated in the tropics as well as in greenhouses in temperate regions. Although local depletion of the resource may occur, the species is not threatened and the diversity still offers opportunities for further selection either for chemical constituents or as an ornamental.

Flame lily shows interesting potential as a source of colchicine, which is highly valued as a medicament for gout. As it is commonly grown as a garden ornamental in the tropics, there might be possibilities for the local production of drugs.

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• N. Bunyapraphatsara & J.L.C.H. van Valkenburg