Euphorbiaceae
Temporal range:
Parts of the candlenut tree
(Aleurites moluccana)
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Malpighiales
Family: Euphorbiaceae
Juss.[1]
Subfamilies
Euphorbia characias flowers

Euphorbiaceae (/juːˈfoʊrbiːˌeɪsiˌaɪ, -siːˌiː/), the spurge family, is a large family of flowering plants. In English, they are also commonly called euphorbias,[2] which is also the name of the type genus of the family. Most spurges, such as Euphorbia paralias, are herbs, but some, especially in the tropics, are shrubs or trees, such as Hevea brasiliensis. Some, such as Euphorbia canariensis,[3]: 206  are succulent and resemble cacti because of convergent evolution.[4] This family has a cosmopolitan global distribution. The greatest diversity of species is in the tropics; however, the Euphorbiaceae also have many species in nontropical areas of all continents except Antarctica.

Description

Cyathia of Euphorbia baylissii
Croton cultivar 'Petra'

The leaves are alternate, seldom opposite, with stipules. They are mainly simple, but where compound, are always palmate, never pinnate. Stipules may be reduced to hairs, glands, or spines, or in succulent species are sometimes absent.

The plants can be monoecious or dioecious. The radially symmetrical flowers are unisexual, with the male and female flowers usually on the same plant. As can be expected from such a large family, a wide variety exists in the structure of the flowers. The stamens (the male organs) number from one to 10 (or even more). The female flowers are hypogynous, that is, with superior ovaries.

The genera in tribe Euphorbieae, subtribe Euphorbiinae (Euphorbia and close relatives) show a highly specialized form of pseudanthium ("false flower" made up of several true flowers) called a cyathium. This is usually a small, cup-like involucre consisting of fused-together bracts and peripheral nectary glands, surrounding a ring of male flowers, each a single stamen. In the middle of the cyathium stands a female flower, a single pistil with branched stigmas. This whole arrangement resembles a single flower.

The fruit is usually a schizocarp, but sometimes a drupe. A typical schizocarp is the regma, a capsular fruit with three or more cells, each of which splits open explosively at maturity, scattering the small seeds.

The family contains a large variety of phytotoxins (toxic substances produced by plants), including diterpene esters, alkaloids, and cyanogenic glycosides (e.g. root tubers of cassava). The seeds of the castor oil plant Ricinus communis contain the highly toxic carbohydrate-binding protein ricin.[5]

A milky latex is a characteristic of the subfamilies Euphorbioideae and Crotonoideae, and the latex of the rubber tree Hevea brasiliensis is the primary source of natural rubber. The latex is poisonous in the Euphorbioideae, but innocuous in the Crotonoideae. White mangrove, also known as blind-your-eye mangrove latex (Excoecaria agallocha), causes blistering on contact and temporary blindness if it contacts the eyes, hence its name. The latex of spurge was used as a laxative.

Twenty first century molecular studies have shown that the enigmatic family Rafflesiaceae, which was only recently recognized to belong to order Malpighiales, is derived from within the Euphorbiaceae.[6]

Euphorbiaceae are monoecious and open pollinated and so self-incompatibility is rare - although it has been reported in the past, apparently this was in error. It is confirmed to be absent or incomplete in herbaceous Chamaesyce by Ehrenfeld 1976, Hevea by Bouharmont 1962, and Manihot by Jennings 1963 and George & Shifriss 1967.[7]

Taxonomy

The family Euphorbiaceae is the fifth-largest flowering plant family[8] and has about 7,500 species[9] organised into 300 genera,[8] 37 tribes, and three subfamilies: Acalyphoideae,[8] Crotonoideae and Euphorbioideae.[10] Amongst the oldest fossils of the group include the permineralised fruit Euphorbiotheca deccanensis from the Intertrappean Beds of India, dating to the late Maastrichtian at the end of the Cretaceous, around 66 million years ago.[11]

Uses and toxicity

Some species of Euphorbiaceae have economic significance, such as cassava (Manihot esculenta), castor oil plant (Ricinus communis), Barbados nut (Jatropha curcas), and the Pará rubber tree (Hevea brasiliensis). Many are grown as ornamental plants, such as poinsettia (Euphorbia pulcherrima) or garden croton (Codiaeum variegatum). Leafy spurge (Euphorbia esula) and Chinese tallow (Triadica sebifera) are invasive weeds in North America.[12]

Seeds of the castor oil plant (Ricinus communis L.) contain the extremely potent toxin, ricin.

Although some species of the Euphorbiaceae have been used in traditional medicine,[13] as of 2019, there is no rigorous clinical evidence that euphorbia extracts are effective for treating any disease.

There is evidence that euphol, a tetracyclic triterpene alcohol, and the main constituent of the sap of the medicinal plant Euphorbia tirucalli, has anti-cancer activity.[14] Analysis of toxicological screening of the inhibitory effect and bioactivity of euphol has shown concentration-dependent cytotoxic effects on cancer cell lines, with more than a five-fold difference in the IC50 values in some cell lines. Euphol treatment had a higher selective cytotoxicity index (0.64-3.36) than temozolomide (0.11-1.13) and reduced both proliferation and cell motility.Euphol also exhibited antitumoral and antiangiogenic activity in vivo, using the chicken chorioallantoic membrane assay, with synergistic temozolomide interactions in most cell lines. In conclusion, euphol exerted in vitro and in vivo cytotoxicity against glioma cells, through several cancer pathways, including the activation of autophagy-associated cell death.[15]

Numerous Euphorbiaceae species are listed on the poisonous plant database of the US Food and Drug Administration mainly because of the toxic sap.[16]

Phytochemistry

Phytochemicals found in Euphorbiaceae species include diterpenoids, terpenoids, flavonoids, alkaloids, tannins, neriifolins (also found in oleander), cycloartenol, lectin, and taraxerol, among others.[13][17]

Conservation

Some species of this family are facing the risk of extinction.[18][19] These include the Euphorbia species E. appariciana, E. attastoma, E. crossadenia,[20] and E. gymnoclada.

References

  1. ↑ Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x.
  2. ↑ "Definition of EUPHORBIA". Merriam Webster Dictionary. Retrieved 2019-06-10.
  3. ↑ Bramwell, D.; Bramwell, Z. (2001). Wild Flowers of the Canary Islands (2nd ed.). Madrid: Rueda. ISBN 978-8472071292.
  4. ↑ Natasha Nguyen (2014). "Convergent evolution of cacti and euphorbias". Retrieved 31 March 2007.
  5. ↑ Wedin GP, Neal JS, Everson GW, Krenzelok EP (May 1986). "Castor bean poisoning". The American Journal of Emergency Medicine. 4 (3): 259–261. doi:10.1016/0735-6757(86)90080-X. PMID 3964368.
  6. ↑ Baum, David A.; Wurdack, Kenneth J.; Nickrent, Daniel L.; Latvis, Maribeth; Davis, Charles C. (2007-03-30). "Floral Gigantism in Rafflesiaceae". Science. 315 (5820): 1812. Bibcode:2007Sci...315.1812D. doi:10.1126/science.1135260. ISSN 0036-8075. PMID 17218493. S2CID 27620205.
  7. ↑ Webster, G. L. (2014). "Euphorbiaceae". In Kubitzki, Klaus (ed.). The Families and Genera of Vascular Plants - Volume XI - Flowering Plants, Eudicots - Malpighiales. Heidelberg: Springer. pp. 51–216/x+331. ISBN 978-3-642-39416-4. OCLC 868922400. ISBN 978-3-642-39417-1. ISBN 3642394167.
  8. 1 2 3 Gillespie, Lynn J.; Armbruster, W. Scott (1997). "A Contribution to the Guianan Flora: Dalechampia, Haematostemon, Omphalea, Pera, Plukenetia, and Tragia (Euphorbiaceae) with Notes on Subfamily Acalyphoideae". Smithsonian Contributions to Botany (86): 6. doi:10.5479/si.0081024X.86. Retrieved 21 May 2018.
  9. ↑ "The Plant list: Euphorbiaceae". Royal Botanic Gardens Edinburgh and Missouri Botanic Gardens. Retrieved 31 March 2017.
  10. ↑ Gurcharan Singh (2004). Plants Systematics: An Integrated Approach. Enfield, N.H: Science Publishers. ISBN 1-57808-351-6.
  11. ↑ Reback, Rachel G.; Kapgate, Dashrath K.; Wurdack, Ken; Manchester, Steven R. (2022-02-01). "Fruits of Euphorbiaceae from the Late Cretaceous Deccan Intertrappean Beds of India". International Journal of Plant Sciences. 183 (2): 128–138. doi:10.1086/717691. ISSN 1058-5893. S2CID 239507275.
  12. ↑ Gucker, Corey L (2010). "Euphorbia esula". Fire Effects Information System, U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory. Archived from the original on 20 May 2022. Retrieved 10 June 2019.
  13. 1 2 Mondal, Sumanta; Ghosh, Debjit (2016). "A complete profile on blind-your-eye mangrove Excoecaria Agallocha L. (Euphorbiaceae): Ethnobotany, phytochemistry, and pharmacological aspects". Pharmacognosy Reviews. 10 (20): 123–138. doi:10.4103/0973-7847.194049. PMC 5214557. PMID 28082796.
  14. ↑ Cruz, L. S.; De Oliveira, T. L.; Kanunfre, C. C.; Paludo, K. S.; Minozzo, B. R.; Prestes, A. P.; Wang, M.; Fernandes, D.; Santos FAD; Manda, V. K.; Khan, S. I.; Ali, Z.; De Messias-Reason, I. J.; Avula, B.; Khan, I. A.; Beltrame, F. L. (2018). "Pharmacokinetics and cytotoxic study of euphol from Euphorbia umbellata (Bruyns) Pax latex". Phytomedicine. 47: 105–112. doi:10.1016/j.phymed.2018.04.055. PMID 30166094. S2CID 52132966.
  15. ↑ Silva VAO; Rosa, M. N.; Miranda-Gonçalves, V.; Costa, A. M.; Tansini, A.; Evangelista, A. F.; Martinho, O.; Carloni, A. C.; Jones, C.; Lima, J. P.; Pianowski, L. F.; Reis, R. M. (2019). "Euphol, a tetracyclic triterpene, from Euphorbia tirucalli induces autophagy and sensitizes temozolomide cytotoxicity on glioblastoma cells". Investigational New Drugs. 37 (2): 223–237. doi:10.1007/s10637-018-0620-y. hdl:1822/57883. PMID 29931585. S2CID 49329684.
  16. ↑ "FDA Poisonous Plant Database". US Food and Drug Administration. 2019. Archived from the original on 15 October 2021. Retrieved 10 June 2019.
  17. ↑ Mali, Prashant Y.; Panchal, Shital S. (2017). "Euphorbia neriifolia L.: Review on botany, ethnomedicinal uses, phytochemistry and biological activities". Asian Pacific Journal of Tropical Medicine. 10 (5): 430–438. doi:10.1016/j.apjtm.2017.05.003. ISSN 1995-7645. PMID 28647179.
  18. ↑ Olson, Mark E.; Lomelí S., José A.; Cacho, N. Ivalú (2005). "Extinction threat in the Pedilanthus clade (Euphorbia, Euphorbiaceae), with special reference to the recently rediscovered E. conzattii (P. pulchellus)". American Journal of Botany. 92 (4): 634–641. doi:10.3732/ajb.92.4.634. ISSN 0002-9122. PMID 21652441.
  19. ↑ Rønsted, Nina; Horn, James W.; Simonsen, Henrik Toft; Nilsson, Niclas; Grace, Olwen M.; Saslis-Lagoudakis, C. Haris; Ernst, Madeleine (2016-07-28). "Evolutionary prediction of medicinal properties in the genus Euphorbia L." Scientific Reports. 6: 30531. Bibcode:2016NatSR...630531E. doi:10.1038/srep30531. ISSN 2045-2322. PMC 4964329. PMID 27464466.
  20. ↑ Zaya, David N.; Howe, Henry F. (2009). "The anomalous Kentucky coVeetree: megafaunal fruit sinking to extinction?". Oecologia. Oecologia: Springer-Verlag. 161 (2): 221–226. Bibcode:2009Oecol.161..221Z. CiteSeerX 10.1.1.536.3841. doi:10.1007/s00442-009-1372-3. PMID 19488788. S2CID 18015.
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