<big>''[[Physalis]]'' L.</big>
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:Protologue: Sp. pl. 1: 182 (1753); Gen. pl. ed. 5: 85 (1754).
== Major species ==
*''Physalis angulata'' L., *''P. minima'' L.
== Vernacular names ==
*Husk tomato, ground cherry (En). *Coqueret (Fr).
== Origin and geographic distribution ==
''Physalis'' consists of about 100 species, mainly distributed in tropical and temperate America, but some of the species have a worldwide distribution. ''P. angulata'' and ''P. minima'' are widely spread as weeds in the Old World, probably as post-Columbian introductions.
== Uses ==
Phytochemically, ''P. angulata'' and ''P. minima'' are well distinguished by the presence of several steroidal lactones, in general belonging to the physaline- and withanolide type. ''P. angulata'' contains the physalins A-I, as well as the withanolides physagulin A-G, withangulatin A and withanolide T, and several vitasteroids (sometimes also mentioned as withasteroids) e.g. vamonolide. ''P. minima'' contains the physalins physalin D, withaphysalin D and E, and dihydroxyphysalin B, and the withanolide withaminimin.
The pyrrolidine alkaloid phygrine (bis-hygrine) was isolated from the roots and aerial parts of both species, as well as from several other ''Physalis'' .
The isolated physalins B and F were found to inhibit the growth of several human leukaemia cells in vitro: K562 (erythroleukaemia), APM1840 (acute T lymphoid leukaemia), HL-60 (acute promyelocytic leukaemia), KG-1 (acute myeloid leukaemia), CTV1 (acute monocytic leukaemia) and B cell (acute B lymphoid leukaemia). In general, physalin F was found to be the most active one. Physalin F also showed cytotoxicity in vitro on 5 other human cancer cell lines, HA22T (hepatoma), HeLa (cervix uteri), KB (nasopharynx), Colo-205 (colon) and Calu-1 (lung), and 3 animal cancer cell lines, H1477 (melanoma), Hep-2 (laryngeal) and 8401 (glioma). The anti-hepatoma and anti-HeLa actions were found to be the strongest. In addition, physalin F had an antitumour effect in mice in vivo, against P388 lymphocytic leukaemia, whereas physalin D was inactive, both in vitro and in vivo.
Isolated withangulatin A, was found to be a topoisomerase II inhibitor in vitro, and a cytotoxic, capable of suppressing general protein synthesis and of inducing the synthesis of a small set of proteins, including those generated by heat-shock treatment, in vivo, in 9L rat brain tumour cells.
Vitanolides, isolated from the aerial parts of ''P. angulata'' , showed anti-inflammatory activity in rat and mice induced inflammation models, although they were less potent than hydrocortisone as reference. Furthermore, the crude leaf extract of ''P. minima'' contains hyperoside (quercetin-3-O-galactoside), which shows a marked anti-inflammatory activity in the rat paw oedema test.
Other biological effects of ''Physalis'' extracts include a marked activity against human African sleeping sickness, caused by ''Trypanosoma brucei rhodesiense'' by petroleum ether-, dichloromethane-, methanol- and water extracts from the aerial parts of ''P. angulata'' , as well as for cytotoxicity for the human fibroblast cell-line WI-38. The tincture of aerial parts macerated in 50% alcohol also showed a marked in vitro inhibitory effect against ''Neisseria gonorrhoeae'' strains isolated from symptomatic patients. The ethanol extract of entire plants showed immunomodulating effects via blastogenesis stimulation in cell cultures. However, immunosuppressive activity was observed when administered intraperitoneally to mice.
Finally, the bright orange colour of several ''Physalis'' fruits is due to the presence of characteristic xanthophylls, e.g. zeaxanthin and cryptoxanthin.
== Adulterations and substitutes ==
Some ''Solanum'' species are used in a similar way as ''Physalis'' to cure digestive and intestinal problems, including stomach-ache and diarrhoea, and for various skin problems such as sores, boils and cuts. Many ''Physalis'' species are also employed to treat fever and malaria, headache and rheumatism. Several other ''Solanaceae'' also contain series of withanolides, e.g. the genera ''Withania'' Pauquy and ''Nicandra'' Adans ''.''
== Description ==
*Annual or perennial herbs, small or large, soft stemmed, angular, often much branched, taproot short. *Leaves alternate to subopposite, simple, usually ovate, sometimes linear, margins toothed to entire, usually hairy, often with trichomes; petiole present; stipules absent. *Flowers axillary, solitary or few together in a fascicle, nodding, actinomorphic, pedicel present; calyx campanulate, 5-toothed; corolla plicate, often open campanulate to rotate, slightly 5-lobed, normally yellowish, often with 5 more or less clear, brownish spots at the throat, varying amounts of hair usually present at the throat; stamens 5, implanted at the base of the corolla tube, anthers 2-celled, opening by a longitudinal slit; style filiform, stigma capitate. *Fruit a biloculed pulpy berry, usually sessile, nodding, enclosed in the persistent, inflated bladdery calyx (husk), 5-10 angled or ribbed. *Seeds numerous, orbicular to reniform, small. *Seedling with epigeal germination.
== Growth and development ==
== Other botanical information ==
''Physalis'' is variable and taxonomically confusing, and no comprehensive study of the genus exists. It is closely related to ''Margaranthus'' and ''Nicandra'' . Intermediate specimens are found between related species, such as ''P. angulata'' and ''P. minima'' in South-East Asia. Both might be varieties of the same species. In America, ''P. angulata'' is morphologically also very similar to ''P. philadelphica'' , of which a larger-fruited, cultivated type and a smaller-fruited, wild type exist.
== Ecology ==
''Physalis'' can be found up to 3000 m altitude at temperatures above 10°C10 °C, although light frost does not kill them. At high temperatures the plants do not develop well. They grow best in moist, fertile soils and are tolerant of partial shade. ''P. angulata'' and ''P. minima'' occur widely as weeds of annual and perennial crops, in waste areas and pastures.
== Propagation and planting ==
In a germination test of ''P. angulata'' seeds, temperatures of 20, 25 and 30°C 30 °C gave about 5, 40 and 45% germination respectively, but at constant temperatures of 10 or 40°C 40 °C no germination occurred. Alternating temperatures for 10 h at 21°C 21 °C and for 14 h at 30°C 30 °C gave about 95% germination. Emergence decreased from about 85% for seeds planted 0-1.2 cm deep to 60% at 5 cm deep, and no seedlings emerged when seeds were planted 10 cm deep. Seeds germinate both in the light and the dark, and germination is optimal when pH is 6-8. Tillage promotes germination slightly compared with no tillage, but germination in both cases is low (about 15% and lower). Irrigation diminishes emergence to less than 5%.
== Diseases and pests ==
''Physalis'' is sensitive to many fungal diseases, caused e.g. by ''Albugo'' , ''Alternaria'' , ''Bipolaris'' , ''Cercospora'' , ''Curvularia'' , ''Odium'' , ''Oidiopsis'' , ''Pseudocercospora'' , ''Pyricularia grisea'' and ''Pythium'' . ''P. angulata'' is also a host of the causal agent of tomato bacterial spot ( ''Xanthomonas campestris'' pv. ''vesicatoria'' ). ''Physalis'' hosts viruses found in tobacco, potato, okra, ''Capsicum'' pepper, beans and several other crops, as well as physalis mottle virus, and also several root-knot nematodes ( ''Meloidogyne'' spp.). ''P. minima'' hosts the parasitic weed ''Orobanche'' ''indica'' Buch.-Ham. ex Roxb.
The inflated calyx of ''Physalis'' protects the fruit-eating larvae of ''Heliothis subflexa'' and ''H. virescens'' ( ''Lepidoptera'' ) (which are also pests on cotton, soyabean and tobacco) against parasitism by the braconid ''Cardiochiles nigriceps'' . ''P. angulata'' is a host for the caterpillar ''Diacrisia obliqua'' , and several plant mites.
== Harvesting ==
== Yield ==
Single plants of ''P. angulata'' may produce 13 100-–31 31 000 seeds. In a field test in Indonesia, the average number of fruits/plant was about 130, and the average number of seeds/fruit was about 130, so about 17 000 seeds per plant were produced.
== Handling after harvest ==
== Literature ==
* Caceres, A., Menendez, H., Mendez, H., Cohobon, E., Samayoa, B.E., Jauregui, E., Peralta, E. & Carrillo, G., 1995. Antigonorrhoeal activity of plants used in Guatemala for the treatment of sexually transmitted diseases. Journal of Ethnopharmacology 48(2): 85-88.
* Waterfall, U.T., 1967. Physalis in Mexico, Central America, and the West Indies. Rhodora 69: 82-120, 203-239, 319-329.
* Young Menzel, M., 1951. The cytotaxonomy and genetics of Physalis. Proceedings of the American Philosophical Society 95(2): 132-183.
== Selection of species ==
<div style="column-count:3;-moz-column-count:3;-webkit-column-count:3">
*[[Physalis angulata (PROSEA)|''Physalis angulata'']]
*[[Physalis minima (PROSEA)|''Physalis minima'']]
*[[ (PROSEA)|'''']]
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== Authors ==
*Slamet Sutanti Budi Rahayu
[[Category:Medicinal plants (PROSEA)]]
[[Category:PROSEA]]