Prota 11(1): Medicinal plants/Plantes médicinales 1
Stirp. nov. 6: 143, pl. 69 (1791).
2n = 36
Endod, soap berry, African soap berry (En). Phytolaque, endod (Fr). Fitolaca, endod (Po).
Origin and geographic distribution
Phytolacca dodecandra is native to sub-Saharan Africa and Madagascar. It has been introduced in Asia and tropical America.
In Central and East Africa and Madagascar various parts of Phytolacca dodecandra are widely used as a medicine for numerous ailments, despite the toxicity of the plant. An extract of the roots, leaves, fruits and seeds is taken as a purgative for humans and animals, as an anthelmintic and to treat oedema and intestinal problems such as diarrhoea and abdominal pain, as a laxative, emetic, sudorific and diuretic. The leaf sap, and sometimes the crushed roots and fruits, are applied to wounds and skin ailments such as ringworm, scabies, eczema, psoriasis, leprosy, boils and vitiligo; the sap is cicatrizing and haemostatic and causes a burning sensation on the skin. In DR Congo the fresh, dried or powdered leaves are used for the same purposes. An infusion of the fruit or roots is taken orally and the young leaves and shoots are chewed to induce abortion. A large lump of butter is taken to stop further contractions. An infusion of the fruit or the root decoction is also widely taken to treat venereal diseases, bilharzia, rabies, malaria, sore throat and other respiratory problems, rheumatic pain, jaundice, as well as anthrax and leeches in animals. In southern Nigeria the leaf decoction is given to newborn babies as a gentle laxative. In Central Africa the mashed leaves are eaten with banana as a stimulant and tonic, especially after childbirth. In Congo the leaf sap is used as eye drops to cure conjunctivitis and river blindness. In Rwanda leaf sap is used as ear drops to treat otitis. In DR Congo the boiled leaves with chicken are given to children with asthma and tuberculosis. In East Africa ground leaves are applied to tumours; the root decoction is also drunk to cause vomiting as treatment of enlarged glands. In Tanzania macerated leaves or root bark are used to treat epilepsy. In Madagascar a decoction of the aerial parts is applied to treat haemorrhoids. In South Africa the root infusion is taken to treat infertility. In Uganda and DR Congo certain cattle tribes use the plant to increase milk production of cows.
The fruits, when dried, powdered and mixed with water, yield a foaming detergent, traditionally used in Ethiopia, Somalia and Uganda for washing clothes, and also to wash the body. Soap has also been made from the ashes of burnt plants in southern Nigeria, or from the fresh leaves in Angola.
In Ethiopia and Zimbabwe unripe fruits are widely applied to control bilharzia-transmitting snails, as they contain molluscicidal saponins. It is a cheaper and less toxic product than synthetic molluscicides. In East Africa the pounded plant is also used as a poison to catch fish.
There is considerable difference in opinion about the edibility of the leaves. In Côte d’Ivoire and southern Nigeria a sauce or soup is prepared from the young shoots and young leaves, and in DR Congo the leaves are cooked as a vegetable. In eastern and southern Africa the whole plant is considered poisonous, and it is said to have caused accidental death of people eating the leaves as a vegetable. In West Africa and Ethiopia the leaves are considered edible for cattle and goats, whereas in most of eastern and southern Africa they are considered poisonous. In Gabon the fruits are eaten. In East Africa the stems are used as ties in the construction of huts and fences. The fruits yield a red dye and the leaves a yellow dye. In Zimbabwe the most common use of the fruits is to colour the floor in houses. In Ethiopia Phytolacca dodecandra is planted as a hedge, and the leaf or fruit extract is sometimes added to drinks and foods as a stimulant, and to curdle milk. The plant is usually not used as firewood, as the smoke is believed to reduce the male sexual ability.
Production and international trade
Small plantations of Phytolacca dodecandra exist in Ethiopia, Zambia, Zimbabwe and Swaziland, and in Ethiopia the fruits are commonly sold in the market for washing clothes.
The leaves, fruits and roots contain numerous saponins (triterpenoid glycosides). These compounds cause haemolysis of red blood cells. The aglycones of the glycosides are mainly composed of oleanolic acid (66%), bayogenin (15%), hederogenin (9%), and 2-hydroxyoleanolic acid (6.5%). On a dry weight basis the fruit pulp contains 25% saponins, and the non-saponin fraction contains a lipid fraction (palmitic acid, oleic acid, stearic acid and a non saponifiable bright orange, waxy material), sugars, starches, pectins and gums and a water insoluble fraction. The saponins have become important as they have strong molluscicidal activity against a range of water snails, and are readily soluble in water and easy to isolate. These properties are useful in the destruction of these snails, including vectors of human diseases such as bilharzia (schistosomiasis), which is caused by parasitic Schistosoma worms. However, the saponins are not active against the egg clusters of the snails. The active saponins have oleanolic acid as the aglycon, the 3 most important being oleanoglycotoxin A, lemmatoxin and lemmatoxin-C. The saponins in the fruit pulp are not molluscicidal, but become active after hydrolytic removal of one of the sugars. The hydrolytic enzyme is contained in the seed and to secure contact between saponins and enzyme and ensure the release of the molluscicidal saponins it is important to finely crush the fruits before soaking them in water. Green unripe fruits contain more active saponins than ripe ones.
Eating fruits or leaves may cause acute poisoning resulting in nausea, bloody diarrhoea, intense congestion of the stomach and intestines, vomiting, weakness, weak and irregular pulse, dilated pupils, swelling of the mucous membrane in the mouth and stupor; death may occur within a few days. Although the solutions of crushed fruits kill a range of aquatic life forms, including small fish, leeches, mosquito larvae, other stages of the bilharzia life-cycle (miracidia and cercariae), zebra mussels and tadpoles, the active ingredients are easily biodegradable and are eliminated from the water within 48 hours. Insect larvae and tadpoles are not affected at the concentrations that kill fish and snails. The molluscicidal potency remains stable over a wide range of pH (5–9), in the presence of various concentrations of organic and inorganic matter and also after irradiation with ultraviolet light. In acute mammalian toxicity tests the fruit extracts were classified as either non-toxic or slightly toxic, except for the eye, in which they can cause severe irritancy. Eye protection is therefore recommended during fruit crushing and handling of dry powders. Ecotoxicity tests indicated that the crushed fruits are no more toxic than currently recommended synthetic molluscicides. Toxicological studies also showed that the fruit extracts do not have mutagenic or carcinogenic properties.
The roots contain saponins of the phytolaccoside or esculentoside types. These compounds showed anti-inflammatory activity. The aqueous extract from the roots showed fungicidal activity against Trichophyton mentagrophytes. Hydroalcoholic extracts of the aerial parts showed significant activity against Staphylococcus aureus, Pseudomonas aeruginosa and Histoplasma capsulatum var. farciminosum, which causes epizootic lymphangitis. The leaf extract showed moderate activity against coxsackie virus in vitro. Butanol extracts of the fruits inhibit the growth of Trichomonas vaginalis, but fermented fruits were inactive. Butanol extracts of the fruits showed spermatocidal properties in vitro and blastocidal activity when injected directly into the uterus of rabbits. Oral administration of a water extract of the aerial parts did not show a significant effect on reproduction in mice.
The leaves and stems produce the antiviral protein dodecandrin, which is a ribosome-inactivating protein (RIP), similar to pokeweed antiviral protein (PAP), isolated from Phytolacca americana L.
Application of the fruit and leaf extract reduced the levels of damage caused by larvae of the maize stalk borer (Busseola fusca), but 2 applications were not sufficient to provide complete protection of maize against second generation larvae.
Studies with callus and cell suspension cultures of Phytolacca dodecandra have established that significant amounts of triterpenoid saponins are produced and retained intracellularly by these cultures. These cultures also produce dodecandrin.
Adulterations and substitutes
Several other plant species contain saponins with molluscicidal activity, including Sapindus saponaria L. and Crossopteryx febrifuga (G.Don) Benth., but the compounds responsible for the activity are often potent fish poisons and harmful to the environment.
Climbing or scrambling dioecious, semi-succulent shrub, sometimes a liana with stems up to 10(–20) m long, with a taproot; trunk sometimes up to 35 cm in diameter; stems usually glabrous. Leaves alternate, simple and entire; stipules absent; petiole 1–4 cm long; blade ovate to broadly elliptical, 3–14 cm × 1.5–9.5 cm, base rounded to slightly decurrent into the petiole, apex acute to rounded, mucronate, glabrous to shortly hairy. Inflorescence an axillary or terminal raceme 5–30 cm long, many-flowered, axis hairy; bracts up to 2.5 mm long, shortly hairy. Flowers functionally unisexual, 5-merous, sweet-scented; pedicel 2–8 mm long; male flowers with narrowly oblong, c. 2.5 mm long, reflexed, whitish to yellowish green sepals, petals absent, stamens 10–20 in 2 whorls, free, filaments 3–7 mm long, ovary usually rudimentary; female flowers with oblong to ovate, c. 2.5 mm long, reflexed sepals, accrescent in fruit, turning yellow to red, petals absent, stamens 8–12, rudimentary, ovary superior, consisting of (4–)5 free, ovoid carpels, styles 1–2 mm long, curved, stigmas linear. Fruit consisting of (4–)5 1-seeded berries fused at base, up to 15 mm in diameter, fleshy, remains of style pointing outwards at apex, ripening orange or purplish red. Seeds kidney-shaped, laterally flattened, 2–4 mm long, shiny black.
Other botanical information
Phytolacca comprises about 25 species, most of which are native to the tropical and subtropical regions of South and Central America, with a few species in Africa and Asia. Phytolacca dodecandra shows extreme morphological variability in leaf hairiness, plant size and growth form.
Another medicinally used Phytolacca species with partly free carpels is Phytolacca heptandra Retz., occurring in Zimbabwe and South Africa. The Xhosa people in South Africa consider the plant very poisonous. A leaf or root extract is commonly drunk as a purgative and emetic. A leaf paste in beer or pulverized root in water is drunk to treat venereal diseases in men. A root decoction is used as an enema to treat enlarged prostate glands. Small amounts of plant pulp are taken as an emetic and applied externally to snakebites. A root maceration is drunk to treat chest pain and delirium. The root is used as bait to kill crows, jackals and stray dogs. In Zimbabwe the plant is also used as a purgative. Lung ailments in cattle are treated by letting them drink a root maceration. The fruit is edible.
Growth and development
In Ethiopia Phytolacca dodecandra flowers and fruits throughout the year, but with a peak during the dry season, from October to April. However, in some parts of Ethiopia it fruits twice a year, in December–February and in June–July. Pollination is effected by insects such as ants, flies and spider wasps. The fruits are often eaten by birds and monkeys, which disperse the seed.
It is reported that the female flowers open later than the male flowers. A well-established plant has only a few but long roots that reach great depths.
Phytolacca dodecandra occurs in forest, forest margins, riparian forest, thickets, wetter bushland, in fences along cultivated land and around houses, on mountain slopes and in open fields, at (0–)1500–3000 m altitude. The plants grow best under direct sunlight in humid, weakly acidic soils that contain high levels of organic matter, in areas with an annual rainfall of about 1400 mm and a distinct dry period. In areas with high evapotranspiration, especially at lower elevations (below 1500 m) partial shade should be available so that the plants do not burn and wilt. Full shade substantially lowers both fruit yield and saponin concentration.
Propagation and planting
Propagation of Phytoloacca dodecandra is by seed or by cuttings. Seed propagation results in equal chances of producing male and female plants, and plants start flowering 2 years after sowing. Germination can be improved by scarification with sand. Fruits can be stored for up to one year without loss of viability, but after 4 years of storage, germination declines by 14%. Propagation by seed is only appropriate for selection purposes. Seed of Phytolacca dodecandra takes about 14 days to germinate. Clonal propagation is possible through non-woody cuttings with 2–3 nodes taken from the top or middle part of the plant. Rooting occurs with or without application of plant growth regulators. Rooted cuttings flower after about 6 months. Tissue culture has been successful for propagation, but the plants take about 18 months to flower. Procedures for routine micropropagation by shoot tip and nodal culture have been developed.
In Ethiopia six-week-old cuttings are transplanted in the field in holes of 60 cm × 60 cm, in a mixture of soil and manure or peat, at the beginning of the rainy season. Survival of transplants depends on availability of water for irrigation during the first and possibly during the second dry season, control of pests, evapotranspiration and soil salinity.
Plantations of Phytolacca dodecandra must be shaded in the first weeks after planting. Occasional watering and weeding are important until the crop has become established. For large-scale cultivation, plants should be spaced 2 m between rows and 1–2 m between plants. For each 10 female plants 1 male plant must be planted to ensure pollination. Annual pruning is necessary to maintain size and shape and to obtain maximum yield. Without pruning the plants become a tangled mass and the fruits are difficult to harvest. Pruning is done by removing the long horizontal branches and shortening erect branches. Intercropping with annual crops is possible because the plant develops few and deep taproots and does not compete much with annual crops for moisture and plant nutrients.
In a field test the best growth and the highest fruit yield were obtained with plants grown in full sunlight, under irrigation and with application of cattle manure. Irrigation was beneficial to fruit yield but reduced the saponin concentration; however, irrigation is beneficial for the total saponin yield. Addition of manure significantly increased growth and fruit yield, but the saponin concentration was lower. The combination of cattle manure and irrigation resulted in the highest total saponin yield.
Diseases and pests
Phytolacca dodecandra is attacked by the larva of different stem- and leafminers (Gitona spp.), which bore through the stem and leaves into the phloem and tunnel in a spiral downwards, leaving a hollow sprout which is easily broken, killing the young shoots. Wilting of young shoots and abortion of inflorescences and fruits are symptoms of infestation. The insects can be chemically controlled by spraying insecticides. Some plants seem to have some natural resistance to the miner; the hairiness of the plant, the presence of raphid crystals in the tissue or the saponin content might contribute to this resistance. Nematodes are also a problem and can infest entire plantations, after which a new plantation site needs to be found.
The fruits of Phytolacca dodecandra are harvested when full grown, but still green, because the saponin content is then highest. Also, ripe fruits are hard to harvest as they fall from the plant and are eaten by birds. Complete fruit bunches are collected manually.
Annual fruit yield increases with the age of the plant up to about 15 years, with the greatest increase occurring between the first and third year.
In Ethiopia several highly molluscicidal and productive cultivars have been selected, and agronomic trials have been effected in eastern and southern Africa. In a field trial in Ethiopia different cultivars yielded between 1050 kg/ha to 2750 kg/ha of dried fruits, with a saponin content of 20–25%. The yield of dry fruits can increase to 3000 kg/ha 4 years after planting, slowly rising to a maximum of about 4000 kg/ha. The content of saponins varies seasonally: fruits harvested during the dry season just before the onset of the rains have the highest content.
Handling after harvest
The harvested fruits must be dried immediately after collection in the open under shade or in the sun. Whole and powdered fruits can be stored at room temperature for up to 4 years without losing potency. Crushed fresh fruits or solutions prepared from them lose their potency within a few days. While grinding the dried fruits care should be taken protect the eyes from the irritating dust.
There are 2 ways of applying the fruits for snail control. The first is to make a slurry of the dried and ground fruits in water. Shortly before application the solution is diluted with river water and administered by siphoning the liquid into the river from a barrel. The rivers are treated when the water level is low and the snails are concentrated in relatively small areas. The second way is to extract the active principles only. This method is suitable for larger water bodies such as lakes, where focal application of the molluscicide is required and sprayers have to be used. In order to avoid blockage of the spray nozzle, an extract has to be prepared. The best results are obtained by soaking the dried and powdered fruits overnight and using butanol to make the extract.
Phytolacca dodecandra is widespread in Africa and is unlikely to be liable to genetic erosion. A major threat is the loss of habitats due to increasing demand for agricultural land. In Ethiopia most populations of Phytolacca dodecandra disappeared during the severe droughts in the 1970s and 1980s. Surviving populations are now almost exclusively confined to protected areas. It is important to preserve the genetic variation of the remaining plants.
In Ethiopia, germplasm collections of Phytolacca dodecandra are maintained at the Institute of Pathobiology and the Ethiopian Plant Genetic Resources Centre in Addis Ababa.
Among 500 different accessions from Ethiopia, 3 strains - E3, E17 and E44 - have been selected for exceptional growth, molluscicidal potency, yield and resistance to insect pests and drought. Based on the colour of the fruits, two types of plants can be distinguished in Ethiopia: ‘Arabe’ with pinkish to red fruits, and ‘Ahiyo’, with yellowish green fruits. The first type has a higher saponin content than the latter one.
A breeding programme needs to be launched to obtain cultivars with high fruit and saponin yields, pest resistance and the ability to grow under the agro-ecological conditions of the lowlands.
Phytolacca dodecandra fruits have great potential as a molluscicide. Agronomic studies to select cultivars with high fruit and saponin yields and pest resistance have been effected, but more research is needed to find ways of making the fruits serve as a supplement or substitute for other detergents. Socio-economic research is needed to evaluate the use of the fruits for local schistosomiasis control.
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Sources of illustration
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Correct citation of this article:
Zimudzi, C., 2007. Phytolacca dodecandra L’Hér. In: Schmelzer, G.H. & Gurib-Fakim, A. (Editors). Prota 11(1): Medicinal plants/Plantes médicinales 1. [CD-Rom]. PROTA, Wageningen, Netherlands.
1, leafy branch with male inflorescence; 2, male flower; 3, female flower; 4, fruit; 5, seed.
Redrawn and adapted by Iskak Syamsudin
leafy branch with young inflorescence
leafy branch with inflorescence
leafy branches with inflorescences
leaves and infructescences