Eclipta prostrata (PROSEA)

Plant Resources of South-East Asia Introduction

List of species

Eclipta prostrata (L.) L.

Protologue: Mant. Pl. 2: 286 (1771).

Family: Compositae

Chromosome number: 2n= 22

Synonyms

• Eclipta alba (L.) Hassk. (1848).

Vernacular names

• False daisy, ink plant (En).
• Herbe à l’encre (Fr)
• Indonesia: orang-aring (Javanese), urang-aring (Sundanese), daun sipat (Moluccas)
• Malaysia: biu, keremak jantan, nigus
• Papua New Guinea: whiteheads (Pidgin)
• Philippines: higis-manok (Tagalog), karim-buaya (Ilokano), pia (Ifugao)
• Laos: hoomz kèèwx
• Thailand: kameng (central), yaa sap, hom kieo (northern)
• Vietnam: cỏ mực, cỏ mhọ nồi, hạn niên thảo, ngô.

Origin and geographic distribution

E. prostrata occurs worldwide in the tropics and subtropics.

Uses

E. prostrata is widely used in traditional medicine against a number of complaints. The aerial parts are generally known for their antibiotic and haemostatic properties and are prescribed for haematuria, nose and wound bleeding and ulcers. The crushed leaves are used against all kinds of skin diseases, especially eczema, but also for mycosis of fingers and feet, and even leprosy, herpes, elephantiasis, haemorrhoids and ringworm. The fresh juice of the plant is applied against scorpion stings and snakebites. The leaves or flower heads are boiled and rubbed on the gums to ease toothache, or used internally for colic, constipation but also against diarrhoea. The root is used in hepatic disorders. In the Philippines, India and Ivory Coast a decoction of the leaves is applied to treat jaundice, while in Indo-China and Brazil the plant is also used against bronchitis and asthma.

In northern Vietnam, stonemasons rub the plant on their hands to stop irritation caused by chalk. The fresh juice of the plant drunk with water is used against tuberculosis. The taste of E. prostrata is sourish, sweet and cooling and the dried leaves, mixed with coconut oil, are used as a tonic. The roots are also known for their purgative and emetic properties, and in Malaysia a decoction of the roots, mixed with cumin, is given to women after childbirth.

In Africa the roots or leaves are used internally for liver and spleen complaints, and for oedema, while in India the juice of the root is applied for conjunctivitis.

The juice from the pounded plant turns black on exposure to air. In Africa, India, Thailand, Indonesia, Malaysia and Indo-China the juice from the fresh or boiled leaves is used for dyeing hair black and encouraging its growth. The sap is used in tattooing; when applied to wounds, it blackens scar-tissue, so that when the wounds are healed the scar is bluish-black.

In Indonesia and Africa the leaves are cooked as a vegetable, and in some parts of India the leaves are used in chutneys.

In Senegal sheep are reported to graze on E. prostrata, while in Kenya it is eaten readily by all cattle.

Production and international trade

In Indonesia, between 1984 and 1990, an estimated 80-120 kg of dried leaves of E. prostrata were traded annually for herbal medicine, both locally and for export.

Properties

E. prostrata contains the coumarin-lactone wedelolactone. Other important constituents include the steroidal alkaloids verazine, eclipalbine and ecliptine, the triterpenoid glucosides ecliptasaponin and ecliptasaponin D, the oleanane triterpene glycosides (echinocystic acid glycosides) eclalbasaponins I-VI, and the taraxastane triterpene glycosides eclalbasaponins VII-X. Also nicotine (0.08%), oleanolic acid, daucosterol, stigmaterol-3-O-glucoside, sitosterol, diosgenin, tigogenin, lanosterol and a number of minerals were isolated from dried E. prostrata plants.

Among the purified isolated compounds of E. prostrata, wedelolactone has been found to be a potent and selective 5-lipoxygenase-inhibitor through an oxygen radical scavenger mechanism with an IC₅₀ of 2.5μM. The steroidal alkaloids all showed weak cytotoxicity against the M-109 cell lines, and in addition, verazine and eclipalbine showed DNA-damaging effects in yeast strains.

The hepatoprotective effects of an extract of E. prostrata were studied against CCl₄- or acetaminophen-induced liver damage in mice and against β-D-galactosamine-induced liver damage in rats. The extract significantly inhibited the acute elevation of serum transaminases induced by CCl₄ and β-D-galactosamine, but had no effect in the acetaminophen model. In another test wedelolactone and desmethylwedelolactone had a significant antihepatotoxic activity employing CCl₄-, β-D-galactosamine-, and phalloidin-induced cytotoxicity in rat hepatocyte cultures. They also showed a significant stimulant effect on liver cell regeneration.

The antimyotoxic and antihaemorrhagic effects of E. prostrata extract, and three of its constituents (wedelolactone, stigmaterol and sitosterol) were investigated in vitro and in vivo. In the models, the myotoxicity of crotalid venoms (Bothrops jararaca, Bothrops jararacussu and Lachesis muta), purified myotoxins (bothropstoxin, bothropasin and crotoxin) and polylysine were quantified in vitro by the release rate of creatine kinase from mouse or rat extensor digitorum muscles, and in vivo by the plasma creatine kinase activity in mice. E. prostrata extract and wedelolactone showed marked antimyotoxic and antihaemorrhagic effects against the crotalid venoms used, which are responsible for most cases of snake poisoning in Brazil. These effects are interpreted as consequences of antiproteolytic and antiphospholipase A2 activities of the constituents.

A decoction of the dried leaves decreased the prothrombin time, it had an anticoumarin activity comparable with vitamin K, and increased the tone of the uterus. In Vietnam, successful results in treating uterine bleeding in pregnant and non-pregnant women have been obtained.

Further biological effects include: antisecretory activity against Escherichia coli, heat-labile and heat-stable enterotoxin-induced secretory responses in rabbit and guinea-pig ileal loop models (extracts of aerial parts), inactivation of hepatitis B surface antigens (hBsAg) in vitro (crude extracts) and induction of mouse skin erythema (extracts of leaves, roots or flower heads).

The leaf extract of E. prostrata shows a marked inhibition of fungal mycelial growth of 8-day-old petri dish colonies of Microsporum gypseum, Trichophyton simii, Malbranchea gypsea or Chrysosporium tropicum. Also, the seeds of E. prostrata in cattle dung show antibacterial activity against Bacillus subtilis, Escherichia coli, Pseudomonas cichorii and Salmonella typhimurium. The alcoholic extract of the whole plant shows antiviral activity against Ranikhet disease virus.

The plant containsα-terthienyl, a sulphur-containing thiophene derivate, which is active against the nematodes Pratylenchus penetrans, Meloidogyne incognita, Hopolaimus indicus, Tylenchus filiformis, Helicotylenchus indicus and Tylenchorhynchus brassicae. It also shows high insecticidal activity against the rice brown plant hopper (Nilaparvata lugens). Finally, E. prostrata extract shows molluscicidal activity at 100 ppm against adult snails and egg masses of Biomphalaria glabrata.

Adulterations and substitutes

Wedelia biflora (L.) DC. leaves or Taraxacum roots are an excellent substitute for roots of E. prostrata in hepatic disorders.

Description

• A straggling annual or rather short-lived perennial herb, (10-)50(-80) cm tall; stem cylindrical, erect or prostrate, rooting at the lower nodes, reddish, with taproot.
• Leaves opposite, simple, oblong-lanceolate, 2-7 cm × 1-2 cm, base attenuate, apex pointed, margins entire or slightly toothed, villous; (sub)sessile; stipules absent.
• Inflorescence an axillary or terminal head, solitary or up to 3 together; peduncle 1-6 cm long, with long, appressed white hairs, involucral bracts 5-6, 2-seriate, oblong, 5-6 mm long, pointed, villous at the back, green, heads ovoid, 5 mm in diameter. The plant has usually numerous, small, white heads.
• Ligulate flowers female, numerous, 2-seriate, up to 3 mm long, apex entire or 2-lobed, white; tubular flowers bisexual, numerous, 4-lobed at the apex, white; anthers 5, obtuse, without appendage at the apex, sagittate at the base; style branches linear, short.
• Fruit an oblong achene, triangular or compressed with 1 central rib on each side, 3 mm × 1.5 mm, truncate at the tip, black; pappus absent, or with 2 weak scales and a few short hairs.
• Seedling with epigeal germination, hypocotyl up to 8.5 mm long, reddish-green, cotyledons ovate, up to 4.5 mm long, base tapering, apex rounded, epicotyl short, densely hairy, first leaves opposite, ovate, margin shallowly toothed, hairy.

Growth and development

E. prostrata is fast growing and early flowering, and may produce more than 17 000 achenes per plant. Plants grown at low temperatures with short day length are shorter and produce many branches, thick leaves and more achenes. Low light intensity increases plant height and leaf size, but decreases specific leaf weight. It is a quantitative short-day plant.

Branching starts from the second week, and is usually finished around the 10th week. Flowering starts from the 5th week, and from the 7th week mature achenes can be found. After the 10th week, achene formation declines.

Other botanical information

Eclipta is a small genus of 3-4 species. Much confusion exists whether the correct name for this taxon is E. alba or E. prostrata, as Linnaeus published both names as Verbesina at the same time in 1753. Later, he published E. erecta (1771), an illegitimate name, because V. alba was cited in synonymy, and also E. prostrata, based on V. prostrata. Hasskarl (1848) united these taxa, and used E. alba (L.) Hassk. as the correct name. Roxburg (1832) however, in his Flora Indica, adopted the name E. prostrata (L.) L., and this name is therefore treated as having priority over E. alba (L.) Hassk.

Ecology

E. prostrata is an anthropogenic species, occurring frequently around houses, open spaces in villages, and disturbed soil. It is a very common weed of rice and sugar cane fields and coconut plantations, and is also found in humid locations along water courses and roadsides, from lowland up to 2000 m altitude. In Sri Lanka, E. prostrata flowers from November to July, in the Philippines all year round. Where rainfall is more than 1200 mm annually, it tends to become perennial. E. prostrata is a polymorphous and troublesome weed in many crops, most difficult in lowland areas with high rainfall. Early control is necessary, and herbicide combinations appear to be more reliable than a single herbicide.

Propagation and planting

E. prostrata propagates through achenes, which are dispersed by water and animals. There is no seed dormancy, so germination can occur throughout the year when moisture is available. When stored at ambient temperatures, the germination rate decreases after about 5 months. In full light, achenes germinate over a range of 10-35°C, but they do not germinate in the dark. Germination is significantly improved by alternating temperatures of 20°C and 35°C. The achenes have a high pH tolerance.

In vitro production of active compounds

Micropropagation of E. prostrata from nodal segment explants was established in a culture of Murashige and Skoog (MS) medium supplemented with 4.4μM benzyladenine. The maximum number of shoots was obtained after 60 days of culture. Wedelolactone was present in shoots cultured in media containing cytokinins, but not in those cultured in basal medium. Callus tissue of E. prostrata grown in an MS culture with D-glucose, shows a considerable increase in ascorbic acid content and growth rate.

Husbandry

E. prostrata is cultivated for biomass and wedelolactone production. Biomass production is not affected by soil pH, but lower pH causes lower concentrations of wedelolactone in the roots.

Diseases and pests

E. prostrata is a host for Macrophomina seolina, powdery mildew in Egypt, and for the mung bean yellow mosaic virus in India. Moreover, it is also a host for the nematodes Meloidogyne incognita, M. javanica, rice root nematode (Hirschmanniella oryzae) and also Trichodorus mirzai. Other surveys indicate that E. prostrata is resistant to Meloidogyne spp., and numerous tests indicate that E. prostrata has strong nematicidal properties against a wide variety of nematodes. The eel-worm (Heterodera radicola) in particular, is attracted by E. prostrata. It would therefore be possible to use E. prostrata to attract and remove this pest from fields with sensitive crops.

E. prostrata is a host for the insects Amsacta moorei, and girdle beetle (Oberea brevis). In field studies conducted in the Philippines, the tephritids Rhabdochaeta asteria and Spathulina acroleuca were observed infesting E. prostrata, and could therefore be potential biological control agents for this weedy species.

Harvesting

In Indo-China, the aerial parts of E. prostrata plants are gathered throughout the year at the end of flowering. In Indonesia, picking of the leaves starts 5-8 weeks after planting, at the beginning of flowering.

Yield

The yield of wedelolactone in vegetative tissues of field-grown E. prostrata plants in Italy is approximately twice that of the roots. Yield of demethylwedelolactone is greater in the roots than in vegetative tissue.

Handling after harvest

In China whole plants of E. prostrata are dried and stored for later use.

Genetic resources and breeding

In Brazil and the United Kingdom, small germplasm collections of E. prostrata exist, and in Indonesia some in vivo collections are present in botanical and medicinal plant gardens. E. prostrata is a pantropical weed, and is not at risk of genetic erosion. No breeding programmes are known to exist.

Prospects

The 5-lipoxygenase inhibition, hepatoprotective, antimyotoxic and antihaemorrhagic effects of E. prostrata extracts, and of its constituents (e.g. wedelolactone), merit further research. Especially the latter two effects might also be of local interest in the treatment of snake bites. In Indonesia, E. prostrata is also a potential plant for the shampoo and the vegetable industry.

Literature

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Authors

• Syamsul Hidayat & S.F.A.J. Horsten