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==Description==
==Description==
''Physalis'' species are [[herbaceous]] plants growing to 0.4 to 3.0 m tall, similar to the common tomato, a plant of the same family, but usually with a stiffer, more upright stem. They can be either annual or [[Perennial plant|perennial]]. Most require full sun and fairly warm to hot temperatures. Some species are sensitive to frost, but others tolerate cold when dormant in winter.
''Physalis'' species are [[herbaceous]] plants growing to {{cvt|0.4|to|3.0|m|ft|0}} tall, similar to the common tomato, a plant of the same family, but usually with a stiffer, more upright stem. They can be either annual or [[Perennial plant|perennial]]. Most require full sun and fairly warm to hot temperatures. Some species are sensitive to frost, but others tolerate cold when dormant in winter.


===Fossil record===
===Fossil record===

Revision as of 12:16, 4 December 2022

"Ground cherry" redirects here. For the European shrub also called "ground cherry", see Prunus fruticosa.

Physalis
Temporal range: Early Eocene (Ypresian) to recent, 52–0 Ma
Cape gooseberry (Physalis peruviana) leaves and fruit
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Solanales
Family: Solanaceae
Subfamily: Solanoideae
Tribe: Physaleae
Genus: Physalis
L.[1]
Type species
Physalis pubescens
Linneaus
Species

About 75-90; see text

Synonyms

Alkekengi Mill.
Herschellia Bowdich ex Rchb.
Pentaphitrum Rchb.[1]

Physalis (/ˈfaɪsəlɪs/, /fɪ-/, /faɪˈseɪlɪs/, /-ˈsæ-/, from φυσαλλίς phusallís "bladder"[2]) is a genus of approximately 75 to 90 flowering plants in the nightshade family (Solanaceae), which are native to the Americas and Australasia. At least 46 species are endemic to Mexico.[3] Cultivated and weedy species have been introduced worldwide. A defining feature of Physalis is a large, papery husk derived from the calyx, which partly or fully encloses the fruit.[4] Many species bear edible fruit, and some species are cultivated.

The typical Physalis fruit is similar to a firm tomato in texture, and like strawberries or pineapple in flavor, with a mild acidity. Some species, such as the Cape gooseberry and tomatillo, have been bred into many cultivars with varying flavors, from tart to sweet to savory. Nations including Colombia, India, and Mexico have a significant economic trade in Physalis fruit.[5] The fruit of many species are generically referred to as physalis, groundcherries,[6] husk tomatoes, husk cherries, poha berries, and golden berries.[7]

Description

Physalis species are herbaceous plants growing to 0.4 to 3.0 m (1 to 10 ft) tall, similar to the common tomato, a plant of the same family, but usually with a stiffer, more upright stem. They can be either annual or perennial. Most require full sun and fairly warm to hot temperatures. Some species are sensitive to frost, but others tolerate cold when dormant in winter.

Fossil record

A 52-million-year-old fossil fruit of Physalis has been found in Patagonia,[8][9] named as Physalis infinemundi.

Cultivation and uses

Physalis peruviana fruit with calyx open

Estimates for the earliest use of Physalis for human consumption range from 900BCE to 5000BCE. Archaeological sites support the historical use of Physalis as a food for indigenous people in what is now northern Mexico and portions of the United States.[5][10][11][12][13]

Physalis fruit are rich in cryptoxanthin. The fruit can be used like the tomato. Once extracted from its husk, it can be eaten raw[14] and used in salads. Some varieties are added to desserts, used as flavoring, made into fruit preserves, or dried and used like raisins. They contain pectin and can be used in pie filling. Ground cherries are called poha in the Hawaiian language, and poha jam and preserves are traditional desserts made from Physalis plants grown on the Hawaiian Islands.[15]

A 2013 literature review identified more than one hundred works with medical use of various Physalis species from the Americas. Preparations included all parts of the plants (fruits, leaves, flowers, stems, and roots) and took forms including decoction, infusion, and soaking. Herbal preparations are known to be administered internally and externally.[16]

Physalis plants grow in most soil types and do very well in poor soils and in pots. They require moisture until fruiting. Plants are susceptible to many of the common tomato diseases and pests, and other pests such as aphids, whiteflies, spider mites, and the false potato beetle (Leptinotarsa juncta) also attack them. Propagation is by seed.

In the United States, Louisiana erroneously classifies Physalis subglabrata (smooth groundcherry) as a hallucinogenic plant, and its cultivation for other than ornamental purposes is outlawed under State Act 159 of 2005. In the Gran Chaco region of South America, the consumption of the different species of Physalis for food has declined due to sociocultural and environmental changes. Factors generally stem from the ongoing effects of colonization, including loss of ancestral territories to forestry exploitation and industrial agriculture as well as the decline of seasonal human migrations which were formerly part of the cycle of propagation, harvest, and consumption of Physalis.[16]

Subgenera and sections

Yellow nightshade groundcherry (Physalis crassifolia)
Physalis peruviana fruits

Physalis is divided into subgenera and sections. The taxonomy of Physalis is still an active area of taxonomic classification.[17] About 75 to 90 species are placed in the genus.[4]

In 1831, Nees von Esenbeck was among the first researchers to complete a review of the Physalis species that had been described until that time in order to identify synonyms and subtaxa within the genus. In 1837, George Don named the sections proposed by Nees von Esenbeck, including three names that are still in use: Physalodendron for woody species, Eurostorhiza for perennials with rhizomatous roots, and Epeteiorhiza for annual species.

Michel Félix Dunal in 1852 and Per Axel Rydberg in 1896 also published efforts to subdivide Physalis, although these were not generally consistent with the taxonomy advocated by Esenbeck and Don. Rydberg's taxonomy identified seven species groups within a section Rydberg called Euphysalis; these groups became the basis of sections which remain in use.[18]

Margaret Y. Menzel's research in the 20th century provided new insights into Physalis taxonomy, including the results of her crossing experiments with 28 Physalis species and their karyological data. The groups of species previously set up by Rydberg were raised to sections by this research. A summary of all taxonomic research regarding the genus was published in 1989 by Radovan Hendrych, and the most recent major taxonomic publications were made in 1994 and 1999 by Mahinda Martínez.[17]

A genetic study by Whitson and Manos in 2005 found evidence that supports the Physalodendron and Rydbergis subgenera. The same research indicated that evidence was weak for most of the recognized species sections within the Rydbergis subgenus, but that other subgroupings might be appropriate instead.[17] This and other phylogenic research led to the Whitson proposal in 2016 to establish Alkekengi officinarum as the type of a new genus rather than the type species of Physalis.[19]

Genetics and breeding

The basic number for Physalis species is 12, and most Physalis species are diploid with 2n = 24. This basic number is typical for members of the Solanaceae family.[20][21][22][23] Research has identified several species that have experienced polyploidy, including Physalis angulata, Physalis floridana, Physalis pubescens, and Physalis peruvania.[24]

Physalis species are generally self-compatible and autogamous,[25] although some may exhibit self-incompatibility and require pollen from another plant to bear fruit or produce seed.[26] A study in 2022 found self-compatibility for all seven Physalis that were observed, which included Physalis peruviana and Physalis ixocarpa. The same study found that fruit fixation and viable seed formation occurred in most inter-specific crosses. The authors did not grow offspring to the second generation.[27]

Menzel performed crosses between Physalis species to assess hybridization in 1951 which showed that perennial species are prone to hybridization while annual Physalis species exhibited barriers between crossing. Hinton identified natural hybridization events between Physalis virginiana Mill. and Physalis heterophylla Nees, which Hinton hypothesized could be the result of self-incompatibility and lack of Physalis virginiana pollen. Sullivan reported in 1985 that natural hybridization rarely occurs among four species from the Physalis viscosa complex.[28]

Physalis subgenus Physalodendron

Authority: (G. Don) M. Martinez

Physalis subgenus Rydbergis

Authority: Hendrych

Section Angulatae

Authority: (Rydberg) M. Martinez

Section Campanulae

Authority: M. Martinez

Section Coztomatae

Authority: M. Martinez

Section Epeteiorhiza

Authority: G. Don

physalis pruinosa plants in bloom

Section Lanceolatae

Authority: (Rydberg) M. Y. Menzel

Section Rydbergae

Authority: M. Martinez

Section Tehuacanae

Authority: M. Martinez

Section Viscosae

Authority: (Rydberg) M. Y. Menzel

Physalis species not assigned to a subgenus or section

Formerly placed here

References

  1. ^ a b "Genus: Physalis L." Germplasm Resources Information Network. United States Department of Agriculture. 2009-09-01. Archived from the original on 2010-05-29. Retrieved 2010-04-14.
  2. ^ "Physalis | Definition of physalis in English by Oxford Dictionaries". Archived from the original on September 22, 2018.
  3. ^ a b c Vargas, O.; et al. (2001). "Two new species of Physalis (Solanaceae) endemic to Jalisco, Mexico". Brittonia. 53 (4): 505–10. doi:10.1007/bf02809650. S2CID 11564.
  4. ^ a b Whitson, M.; Manos, P. S. (2005). "Untangling Physalis (Solanaceae) from the physaloids: a two-gene phylogeny of the Physalinae". Systematic Botany. 30 (1): 216–30. doi:10.1600/0363644053661841. JSTOR 25064051. S2CID 86411770.
  5. ^ a b Vargas-Ponce, Ofelia; Sánchez Martínez, José; Zamora Tavares, María del Pilar; Valdivia Mares, Luis Enrique (2016-12-01). "Traditional management of a small-scale crop of Physalis angulata in Western Mexico". Genetic Resources and Crop Evolution. 63 (8): 1383–1395. doi:10.1007/s10722-015-0326-3. ISSN 1573-5109. S2CID 41108248.
  6. ^ "Physalis". Integrated Taxonomic Information System. Retrieved 2011-05-21.
  7. ^ Doctor, Vikram (4 March 2013). "Golden berry: Decoding the acid freshness and wild sweet taste of physalis". The Economic Times. Retrieved 6 Sep 2014.
  8. ^ bbc.com/news/science-environment-38511034
  9. ^ Wilf, Peter (6 Jan 2017). "Eocene lantern fruits from Gondwanan Patagonia and the early origins of Solanaceae". Science. 355 (6320): 71–75. Bibcode:2017Sci...355...71W. doi:10.1126/science.aag2737. PMID 28059765. S2CID 206651318.
  10. ^ Jennings, Jesse D. (April 1970). "The Prehistory of the Tehuacan Valley, Vol. 1: Environment and Subsistence. Edited by Douglas S. Byers. University of Texas Press, Austin, 1967. - The Prehistory of the Tehuacan Valley, Vol. 2: The Non-Ceramic Artifacts. Richard S. MACNEISH, Antoinette Nelken-Terner, and Irmgard W. Johnson. University of Texas, Austin, 1967". American Antiquity. 35 (2): 234–236. doi:10.2307/278167. ISSN 0002-7316. JSTOR 278167.
  11. ^ Ruz Sosa, Mario Humberto (2013-02-15). "El añil en el Yucatán del siglo XVI". Estudios de Cultura Maya. 12. doi:10.19130/iifl.ecm.1979.12.525. ISSN 2448-5179.
  12. ^ Le Paige, Gustavo (1977). "Recientes descubrimientos arqueológicos en la zona de San Pedro de Atacama". Estudios Atacameños (5): 111–126. doi:10.22199/s07181043.1977.0005.00008. ISSN 0716-0925.
  13. ^ McClung de Tapia, Emily; Martínez Yrízar, Diana; Ibarra Morales, Emilio; Adriano Morán, Carmen Cristina (January 2014). "Los orígenes prehispánicos de una tradición alimentaria en la cuenca de méxico". Anales de Antropología. 48 (1): 97–121. doi:10.1016/s0185-1225(14)70491-6. ISSN 0185-1225.
  14. ^ Angier, Bradford (1974). Field Guide to Edible Wild Plants. Harrisburg, PA: Stackpole Books. p. 90. ISBN 0-8117-0616-8. OCLC 799792.
  15. ^ Gibbons, Euell (1962). Stalking the Wild Asparagus. Chambersburg, Pennsylvania: Alan C. Hood & Company, Inc. p. 104. ISBN 978-0-911469-03-5.
  16. ^ a b Arenas, Pastor; Kamienkowski, Nicolás Martín (December 2013). "Ethnobotany of the Genus Physalis L. (Solanaceae) in the South American Gran Chaco". Candollea. 68 (2): 251–266. doi:10.15553/c2012v682a9. ISSN 0373-2967.
  17. ^ a b c Whitson, Maggie; Manos, Paul S. (2005). "Untangling Physalis (Solanaceae) from the Physaloids: A Two-Gene Phylogeny of the Physalinae". Systematic Botany. 30 (1): 216–230. doi:10.1600/0363644053661841. ISSN 0363-6445. JSTOR 25064051. S2CID 86411770.
  18. ^ Solanaceae IV : advances in biology and utilization. Michael Nee, International Solanaceae Congress. [Richmond, England]: Royal Botanic Gardens, Kew. 1999. ISBN 1-900347-90-3. OCLC 44910181.{{cite book}}: CS1 maint: others (link)
  19. ^ Whitson, Maggie (April 2011). "(2016) Proposal to conserve the name Physalis ( Solanaceae ) with a conserved type". Taxon. 60 (2): 608–609. doi:10.1002/tax.602047.
  20. ^ Lu, Jiangjie; Luo, Meifang; Wang, Li; Li, Kunpeng; Yu, Yongyi; Yang, Weifei; Gong, Pichang; Gao, Huihui; Li, Qiaoru; Zhao, Jing; Wu, Lanfeng (December 2021). "The Physalis floridana genome provides insights into the biochemical and morphological evolution of Physalis fruits". Horticulture Research. 8 (1): 244. doi:10.1038/s41438-021-00705-w. ISSN 2662-6810. PMC 8602270. PMID 34795210.
  21. ^ Azeez, Sekinat Okikiola; Faluyi, Julius Olaoye; Oziegbe, Mathew (2019-11-08). "Cytological, foliar epidermal and pollen grain studies in relation to ploidy levels in four species of Physalis L. (Solanaceae) from Nigeria". International Journal of Biological and Chemical Sciences. 13 (4): 1960–1968. doi:10.4314/ijbcs.v13i4.4. ISSN 1997-342X. S2CID 209597935.
  22. ^ Escobar-Guzmán, Rocío; Ochoa-Alejo, Neftalí (2021), Segui-Simarro, Jose M. (ed.), "Anther Culture of the Gametophytic Self-Incompatible Species Physalis ixocarpa Brot", Doubled Haploid Technology, vol. 2288, New York, NY: Springer US, pp. 319–326, doi:10.1007/978-1-0716-1335-1_20, ISBN 978-1-0716-1334-4, PMID 34270021, S2CID 235961279, retrieved 2022-08-07
  23. ^ Venkateswarlu, J.; Rao, K. G. Raja (1979). "Morphology of the Pachytene Chromosomes of Physalis pubescens L." Cytologia. 44 (1): 161–166. doi:10.1508/cytologia.44.161. S2CID 84930570.
  24. ^ Liberato, Sara; Sánchez-Betancourt, Erika; Argüelles Cárdenas, Jorge Humberto; Gonzólez, Carolina; Núñez, Víctor (2015-12-30). "Citogenética de genotipos de uchuva, Physalis peruviana L., y Physalis floridana Rydb., con respuesta diferencial a Fusarium oxysporum". Ciencia & Tecnología Agropecuaria. 15 (1): 51–61. doi:10.21930/rcta.vol15_num1_art:396. ISSN 2500-5308.
  25. ^ Azeez, Sekinat Okikiola; Faluyi, Julius Olaoye (2018-06-29). "Hybridization in Four Nigerian Physalis (Linn.) Species". Notulae Scientia Biologicae. 10 (2): 205–210. doi:10.15835/nsb10210241. ISSN 2067-3264. S2CID 90292809.
  26. ^ Peña-Lomelí, Aureliano; Magaña-Lira, Natanael; Gámez-Torres, Adrián; Mendoza-Celino, Fredy Ángel; Pérez-Grajales, Mario (2018). "Polinización manual en dos variedades de tomate de cáscara (Physalis ixocarpa Brot. ex Horm.) en invernadero". Revista Chapingo Serie Horticultura. 24 (1). doi:10.5154/r.rchsh.2017.02.011. ISSN 1027-152X.
  27. ^ Junior, André Dutra Silva; Zeist, André Ricardo; da Silva, Daniel Fernandes; de Souza Leal, Murilo Henrique; Oliveira, Guilherme José Almeida; de Oliveira, Joana Nascimento Merces; Youssef, Khamis; Da Rocha Toroco, Bruno (2022-05-18). "Reproductive biology and hybridization of Physalis L. species". Brazilian Journal of Botany. 45 (3): 1037–1045. doi:10.1007/s40415-022-00811-6. S2CID 248902866 – via Springer Link.
  28. ^ Pretz, Chelsea; Deanna, Rocío (February 2020). "Typifications and nomenclatural notes in Physalis (Solanaceae) from the United States". Taxon. 69 (1): 170–192. doi:10.1002/tax.12159. ISSN 0040-0262. S2CID 214125933.
  29. ^ a b c d e Pretz, Chelsea; Deanna, Rocío (February 2020). "Typifications and nomenclatural notes in Physalis (Solanaceae) from the United States". Taxon. 69 (1): 170–192. doi:10.1002/tax.12159. ISSN 0040-0262. S2CID 214125933.
  30. ^ Pyne, Milo; Orzell, Steve L.; Bridges, Edwin L.; Poindexter, Derick (2019). "Physalis Macrosperma (Solanaceae: Physalinae), A New Psammophyte Endemic to the West Gulf Coastal Plain of the Southeastern U.S.A., A Global Biodiversity Hotspot". Journal of the Botanical Research Institute of Texas. 13 (1): 31–50. doi:10.17348/jbrit.v13.i1.824. ISSN 1934-5259. JSTOR 26783957. S2CID 244520017.
  31. ^ Switek, Brian. "Paleo Profile: Tomatillo from the End of the World".
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