PROTA homepage Prota 11(1): Medicinal plants/Plantes m้dicinales 1
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Solanum aculeatissimum Jacq.

Protologue
Icon. pl. rar. 1: 5, pl. 41 (1781).
Family
Solanaceae
Chromosome number
2n = 24
Synonyms
Solanum khasianum C.B.Clarke (1883), Solanum angustispinosum De Wild. (1914).
Vernacular names
Cockroach berry, love apple, devil’s apple, soda-apple nightshade (En). Pomme d’amour (Fr). Unisca, jurubeba, gogoia (Po).
Origin and geographic distribution
Solanum aculeatissimum is native of Brazil. It was introduced in Africa centuries ago and is now found throughout tropical Africa, but not yet in the Indian Ocean Islands. In Asia it is widely spread as well.
Uses
In Liberia and Nigeria a fruit decoction of Solanum aculeatissimum is administered as an enema to cure constipation. In DR Congo warm leaves are ground in water and the extract is administered as an enema as an abortifacient or purgative. In Uganda the fruit sap is used as eye drops to treat trachoma and the sap of roasted fruits is taken to induce labour. The fruits are considered toxic and the plants are used in witchcraft. In Kenya the sap from the leaves and fruit is applied to warts. In South Africa a root decoction is drunk to cure back pain and male impotence, a root maceration is drunk to treat snakebites, smoke from burning fruits is inhaled to cure toothache and the fruit is pressed to the forehead to remedy headache and to the skin to cure skin infections. The root extract is taken as a purgative and also to stop flatulence. Roots and fruits enter in preparations to treat coughs and dysmenorrhoea. In China the plant is used to treat bronchitis and rheumatism. No cases of poisoning of free ranging cattle are reported but force-feeding of either ripe or unripe fruits leads to death. In Puerto Rico the sliced fruit is used as bait for cockroaches.
In Asia Solanum aculeatissimum is used as rootstock for tomato and eggplant.
Properties
Solasonine is the major glycoalkaloid in leaves, stems, fruits and seeds of Solanum aculeatissimum, and minor alkaloids isolated are solamargine, solanine and solasodine. The highest concentration of alkaloids is found in the seed (4.4%). Ripe fruits have a solasodine content of 2.8–3.8%. Mature fruits contain the steroids diosgenin, β-sitosterol and lanosterol. Roots contain the steroidal saponins aculeatiside A and B, which are precursors of steroid hormones. The roots contain high amounts of solasodine and aculeatiside B (3.8% and 3.0% respectively).
An ethanol extract of the leaves inhibits spore germination of the fungal cabbage disease black spot (Alternaria brassicicola). Extracts of leaves and fruits were shown to have a repellent, but not an insecticidal effect on the cabbage aphid (Brevicoryne brassicae).
In-vitro production of the steroidal saponins is of economic interest; it has been extensively investigated and protocols have been developed.
Description
Perennial herb or small shrub up to 120(–200) m tall; stems densely covered with yellowish, slender prickles up to 18 mm long, hairs mostly simple, some hairs stellate. Leaves alternate, simple; stipules absent; petiole 1–9.5 cm long; blade broadly ovate, 2.5–16 cm ื 2.5–19 cm, base unequal, cuneate to cordate, apex usually acute, margin coarsely toothed or more or less deeply lobed, prickles present on main veins. Inflorescence a cyme, inserted above the leaf axil, 2–3.5 cm long, 2–7-flowered. Flowers bisexual or functionally male, regular, 5(–6)-merous; calyx bell- or cup-shaped, 4–10 mm in diameter, lobes unequal, triangular, acuminate, reflexed; corolla stellate, up to 22 mm in diameter, white to mauve or purple; stamens alternate with corolla lobes, filaments 1–2 mm long, anthers lanceolate, c. 6 mm long, opening with apical pores; ovary superior, with stalked glands and short hairs near base, style glabrous, 6–10 mm long. Fruit a globose berry 2–3 cm in diameter, glabrous, green marbled white or cream, yellowish when ripe, many-seeded. Seeds ovoid, compressed, 2.5–4.5 mm in diameter, margin thickened. Seedling with epigeal germination; cotyledons thin, leafy.
Other botanical information
Solanum comprises about 1000 species and has a cosmopolitan distribution, except in boreal, alpine and aquatic habitats. About 110 species are found in tropical Africa. The principal centre of diversity is located in Central and South America, with secondary centres in Africa and Australia. Solanum has been subdivided into 7 subgenera and numerous sections and series. The section Acanthophora of the subgenus Leptostemonum, to which Solanum aculeatissimum belongs, comprises about 20 species, all native to tropical America. The name Solanum aculeatissimum is sometimes wrongly applied to Solanum capsicoides All., a species native of coastal Brazil. Solanum capsicoides is fairly widespread in the Caribbean and the fruits are used to make juice. It is also introduced in tropical Africa as an ornamental. The synonym Solanum khasianum is still often used in scientific publications in India, and as such it is often misapplied to Solanum viarum Dunal, another species of the section Acanthophora. Based on their morphology it has been concluded that Solanum aculeatissimum, Solanum viarum and Solanum myriacanthum Dunal are very closely related and molecular research indicates that they belong to a hybrid complex. Misidentification of Solanum macrocarpon L. (synonym: Solanum dasyphyllum Schumach. & Thonn.) is a further source of confusion, probably in West Africa in particular.
Growth and development
In East Africa Solanum aculeatissimum flowers from October to February, in DR Congo throughout the year and in South Africa from November to March.
Ecology
Solanum aculeatissimum occurs in disturbed open vegetation, e.g. in roadsides, forest clearings and as a weed in pastures, agricultural land and forest plantations, from sea-level up to 2400 m altitude.
Propagation and planting
Solanum aculeatissimum is easily propagated from seed or stem cuttings. Shoots can be produced from leaf and stem-node explants on Murashige & Skoog medium supplemented with cytokinins.
Management
Commercial cultivation of Solanum aculeatissimum has not been attempted in tropical Africa. In Asia it is used as rootstock for tomato and eggplant, both in traditional systems and in hydroponics.
Diseases and pests
Solanum aculeatissimum is a host of the potato leaf rolling virus (PLRV). It is very susceptible to root knot nematodes (Meloidogyne spp.).
Harvesting
When grown in Japan as an arable crop the dry weight of the roots reached a maximum in November and contained more than 10% steroid saponins.
Genetic resources
In Brazil the genetic diversity of Solanum aculeatissimum should be looked into, with regard to the alkaloid content. Plants grown in botanic gardens in Europe are very uniform and probably descend from a single introduction from South Africa.
Breeding
Solanum aculeatissimum is highly resistant to Verticillium wilt (Verticillium dahliae), a characteristic that is useful to introduce in eggplant (Solanum melongena L.) and gboma eggplant (Solanum macrocarpon L.); crosses of Solanum aculeatissimum with these 2 species are sometimes successful, but sometimes result in sterile seed.
Prospects
The prospects of cultivating Solanum aculeatissimum for the extraction of steroidal saponins need further research. A major problem in evaluating the potential of Solanum aculeatissimum is the enormous confusion in the literature with several other Solanum species.
Major references
• 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.
• Gbile, Z.O. & Adesina, S.K., 1988. Nigerian Solanum species of economic importance. Annals of the Missouri Botanical Garden 75: 862–865.
• Gon็alves, 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.
• Hulstaert, G., 1966. Notes de Botanique Mongo. Acad้mie Royale des Sciences d’Outre-mer, Classe des Sciences Naturelles et M้dicales, N.S. 15–3, Bruxelles, Belgium. 213 pp.
• Levin, R.A., Watson, K. & Bohs, L., 2005. A four-gene study of evolutionary relationships in Solanum section Acanthophora. American Journal of Botany 92(4): 603–612.
• Nabeta, K., 1993. Solanum aculeatissimum Jacq: in vitro culture and the production of secondary metabolites. In: Bajaj, Y.P.S. (Editor). Biotechnology in agriculture and forestry. Volume 24. Medicinal and aromatic plants V. Springer-Verlag, Berlin, Germany. pp. 329–341.
• Nee, M., 1991. Synopsis of Solanum Section Acanthophora. 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. 257–266.
• 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.
• Welman, W.G., 2003. The genus Solanum (Solanaceae) in southern Africa: subgenus Leptostemonum, the introduced sections Acanthophora and Torva. Bothalia 33(1): 1–18.
Other references
• Alconero, R., Robinson, R.W., Dicklow, B. & Shail, J., 1988. Verticillium wilt resistance in eggplant, related Solanum species, and interspecific hybrids. HortScience 23(2): 388–390.
• Bukenya-Ziraba, R. & Carasco, J.F., 1999. Ethnobotanical aspects of Solanum L. (Solanaceae) in Uganda. In: Nee, M., Symon, D.E., Lester, R.N. & Jessop, J.P. (Editors). Solanaceae 4: Advances in biology and utilization. Royal Botanic Gardens, Kew, Richmond, United Kingdom. pp. 345–360.
• 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.
• Heine, H., 1963. Solanaceae. In: Hepper, F.N. (Editor). Flora of West Tropical Africa. Volume 2. 2nd Edition. Crown Agents for Oversea Governments and Administrations, London, United Kingdom. pp. 325–335.
• Hsieh, T.F., Huang, J.H., Hsieh, L.J., Hu, M.F. & Ko, W.H., 2005. Antifungal effect of plant extracts on phytopathogenic fungi. Plant Pathology Bulletin 14(1): 59–66.
• Hutchings, A., Haxton Scott, A., Lewis, G. & Cunningham, A., 1996. Zulu medicinal plants: an inventory. University of Natal Press, Pietermaritzburg, South Africa. 450 pp.
• Ikenaga, T., Kikuta, S., Itimura, K., Nakashima, K. & Matsubara, T., 1988. Growth and production of steroid saponin in Solanum aculeatissimum during one vegetation period. Planta Medica 54: 140–142.
• Kadkade, P.G., 1984. Influence of cultural conditions on glycoalkaloid formation in Solanum aculeatissimum tissue cultures. In: Proceedings of the 11th annual meeting, Plant Growth Regulator Society of America, Boston, Massachusetts, United States. pp. 79–86.
• Kadkade, P.G., Recinos, J.A. & Madrid, T.R., 1979. Studies on the distribution of glycoalkaloids in Solanum aculeatissimum. Planta Medica 37(1): 70–72.
• Kamatenesi-Mugisha, M. & Oryem-Origa, H., 2007. Medicinal plants used to induce labour during childbirth in western Uganda. Journal of Ethnopharmacology 109: 1–9.
• Lovatto, P.B., Goetze, M. & Thom้, G.C.H., 2004. Efeito de extratos de plantas silvestres da familia Solanaceae sobre o controle de Brevicoryne brassicae em couve (Brassica oleracea var. acephala). Ci๊ncia Rural 34(4): 971–978.
• Rathore, A.K. & Kamal, R., 1979. Steroids and steroidal alkaloids of Solanum aculeatissimum. Pharmazie 34(4): 250–251.
• Shale, T.L., Stirk, W.A. & van Staden, J., 1999. Screening of medicinal plants used in Lesotho for anti-bacterial and anti-inflammatory activity. Journal of Ethnopharmacology 67: 347–354.
• De Souza-Dias, J.C., Costa-Alvaro, S. & Nardin-Antonio, M., 1993. Potato leafroll virus in solanaceous weeds in Brazil explains severe outbreaks of the disease in absence of known potato donor sources. Summa Phytopathologica 19(2): 80–85.
• Tokarnia, C.H., Canella, C.F.C. & Dobereiner, J., 1973. Intoxicacao experimental em bovinos pelos frutos de Solanum aculeatissimum. Pesquisa Agropecuaria Brasileira Veterinaria 8(6): 35–39.
• Asano, N., Kato, A., Matsui, K., Watson, A.A., Nash, R.J., Molyneux, R.J., Hackett, L., Topping, J. & Winchester, B., 1997. The effects of calystegines isolated from edible fruits and vegetables on mammalian liver glycosidases. Glycobiology 7(8): 1085–1088.
• Yamada, T., 1999. A report of the ethnobotany of the Nyindu in the eastern part of the former Zaire. African Study Monographs 20(1): 1–72.
Sources of illustration
• Agnew, A.D.Q. & Agnew, S., 1994. Upland Kenya wild flowers: a flora of the ferns and herbaceous flowering plants of upland Kenya. 2nd Edition. East Africa Natural History Society, Nairobi, Kenya. 374 pp.
• Welman, W.G., 2003. The genus Solanum (Solanaceae) in southern Africa: subgenus Leptostemonum, the introduced sections Acanthophora and Torva. Bothalia 33(1): 1–18.
Author(s)
• M.J. Nicholson
Plants for Life, P.O. Box 617, Limuru, Kenya


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, R้duit, 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

Correct citation of this article:
Nicholson, M.J., 2008. Solanum aculeatissimum Jacq. In: Schmelzer, G.H. & Gurib-Fakim, A. (Editors). Prota 11(1): Medicinal plants/Plantes m้dicinales 1. [CD-Rom]. PROTA, Wageningen, Netherlands.
Distribution Map naturalized


1, flowering shoot; 2, fruits.
Redrawn and adapted by Achmad Satiri Nurhaman