View source for Solanum lycocarpum

{{Short description|Species of flowering plant}}
{{Distinguish|Solanum lycopersicum|wolfberry|Wolf River (apple)}}
{{Speciesbox 
|image = Ramo de lobeiro.JPG
|genus = Solanum
|species = lycocarpum
|authority = [[Augustin Saint-Hilaire|A.St.-Hil.]]
}}

'''''Solanum lycocarpum''''', or '''wolf apple,''' is common in the Brazilian [[Cerrado]] [[ecoregion]]. The plant is called '''''lobeira''''' ("wolf's plant") or '''''fruta-do-lobo''''' ("wolf's fruit") in Portuguese.<ref name=motta1996/>

The name "wolf apple" comes from the fact that they account for more than 50% of [[maned wolf|maned wolves']] diet.<ref name=motta1996/> Likewise, the scientific name "''lycocarpum''" is formed from [[Latin]]ized [[Greek language|Greek]] elements "''lyco-''", meaning "wolf", and "''carpum''" meaning "fruit".<ref name=jaeg1944/>

==Range==
The native range of the wolf apple tree is the [[Cerrado|Brazilian savannah]], but it grows also on pastures and disturbed land, such as highway margins, in various parts of Brazil.<ref name=cler2011/> It prefers moist, clay soil, full sun, and mild temperatures.<ref name=oliv1988/><ref name=tata2006/>

==Description==
[[Image:Solanum lycocarpum flower.jpg|left|thumb|Flower]]
[[File:Flickr - João de Deus Medeiros - Solanum lycocarpum (1).jpg|left|thumb|Fruit]]
The wolf apple plant is a flowering [[shrub]] or small tree with round open crown, ranging in height from {{cvt|1.2|to|5|m|ft|0}}.  The large leaves are {{cvt|16|to|28|cm|in|0}} long, simple but deeply lobed, tough, and covered in a soft grey-white fuzz, in alternate disposition.

Plants flower through the year, but more intensely during the dry season, from late fall to early spring. The flowers are similar to those of the [[bitter nightshade]]: star-shaped, with five [[sepal]]s, five blue or purple [[petal]]s fused at the base, and five large yellow [[anther]]s that release [[pollen]] through an opening at the tip.  The flowers are arranged in helical [[inflorescence#Determinate or cymose|monochasal cyme]], opening from the base up. Flowers lack [[nectar]], and are either male or hermaphroditic.  The latter produce fruit after cross-pollination by several species of pollen-collecting [[bee]]s, including ''[[Xylocopa virescens]]'', ''[[Xylocopa frontalis|X. frontalis]]'', ''[[Oxaea flavescens]]'', ''[[Centris scopipes]]'', ''[[Centris fuscata|C. fuscata]]'', ''[[Bombus morio]]'', ''[[Exomalopsis]]'' sp., ''[[Pseudaugochloropsis graminea]]'', ''[[Apis mellifera]]'', and  ''[[Megachile benigna]]''.<ref name=oliv1988/><ref name=tata2006/>

Fruit production occurs mostly between December and January. The fruits are large, up to {{cvt|20|cm|in|0}} in diameter and weighing {{cvt|900|g|lb|0}} or more, and contain up to 500 dark brown comma-shaped seeds, about {{cvt|7|×|5|×|2|mm|in|frac=16}}.<ref name=pint2007/>  The rind is thin and lustrous, and remains green even after ripening.<ref name=pint2007/> The pulp is yellowish, soft, sweetish and extremely aromatic.<ref name=cler2011/>  They resemble the [[tomato]] in external shape and internal morphology,<ref name=tata2006/> but also the [[eggplant]] for the texture and color of the flesh. The maned wolf is an important seed-spreading agent.<ref name=lomb1993/>

==Uses==
===Culinary use===
The ripe fruits are edible to [[humans]], and are consumed by local populations in [[Fruit preserves|jams and preserves]], but the unripe fruits are quite rich in [[tannin]].<ref name=tata2006/> The fruits are used as a fallback source of food for [[cattle]] in the dry season.

At the onset of maturation, the whole fruit contains 85% moisture (by weight), 5% starch, and 10% other residue.<ref name=cler2011/> The dry pulp, without peel and seeds, yields over 50% starch, more than [[cassava]].<ref name=pasc2013/>  The starch is locally separated by blending the fruit with water and straining.<ref name=cler2011/>

The pulp of the ripe fruit contains about 75% moisture, 11% [[carbohydrate]], 1% [[fat]], 1% [[protein]], 1% ash, 2.5% [[fiber (food)|soluble fiber]], and 2% insoluble fiber.  The main volatiles in the ripe fruit are [[hexanal]] (which has the scent of grass) and [[ethyl butanoate]] (scent of apple), although dozens of other compounds contribute to the fruit's aroma.<ref name=pere2019/>

All other parts of the plant are [[poison]]ous to humans.  However, the leaves are eaten by [[cattle]], and are also an important food source for the [[bat]] ''[[Platyrrhinus lineatus]]''.<ref name=agui2005/>

===Medicine===
Currently, there are no proven medicinal uses for the wolf apple, but the plant is used in the local [[folk medicine]] for [[obesity]], cholesterol reduction, and as a treatment for [[diabetes]].<ref name=tata2006/> About 30% of the starch extracted from almost-ripe fruits, which commercialized as capsules, is digestion-resistant,<ref name=cler2011/> which seems to be the rationale for these popular uses.<ref name=agno2000/> High-fiber ''Solanum lycocarpum'' flour was found to reduce diabetes symptoms in diabetic rats;<ref name=fari2010/> however, the starch did not show such effect.<ref name=oliv2003/>

The fruit contains about 1.5% of [[glycoalkaloid]]s in its dried fruits, chiefly [[solamargine]] and [[solasonine]],<ref name=moti1970/> mostly in the peel and seeds.<ref name=cler2011/> Extracts have been shown to kill [[in vitro]] the human parasites ''[[Schistosoma mansoni]]''<ref name=mira2012/> and ''[[Giardia lamblia]]'',<ref name=mart2015/> ''[[Leishmania amazonensis]]'',<ref name=mira2013/> and ''[[Leishmania infantum]]''.<ref name=clem2018/> The main [[phenolic compounds]] in [[ethanol]] extracts of the ripe fruits are [[caffeic acid]] and [[chlorogenic acid]].<ref name=mora2015/>

It is believed that the fruits protect the [[maned wolf]] from infection of the [[Dioctophyme renale|giant kidney worm]], which is usually fatal for the animal.<ref name=tata2006/>

''Solanum lycocarpum'' fruit extracts have also been shown to [[larvicide|kill larvae]] of the [[Culex quinquefasciatus|''Culex'']] [[mosquito]].<ref name=silv2015/>

==Pests==
The wolf apple tree is the natural host of the larva of the [[butterfly]] ''[[Leucanella memusae]]''.  Contact with the urticating bristles of this caterpillar may result in severe [[dermatitis]] and [[hemorrhage]].<ref name=soar2019/>

The [[weevil]] ''[[Collabismus clitellae]]'' lays eggs into stems of the plant, causing the formation of [[gall]]s, with measurable harmful effect on the plant's growth and fruit production.<ref name=souz1998/>

==References==
<references>
<ref name=tata2006>Fernando Tatagiba (2006): "[http://www.biologo.com.br/plantas/cerrado/lobeira.html Lobeira, fruta-de-lobo]".  Online article on the ''Biólogo'' website, published 2006-10-06. Accessed on 2020-06-30.</ref>

<ref name=motta1996>J. C. Motta-Junior, S. A.Talamoni, J. A. Lombardi, and K. Simokomaki (1996): "Diet of the maned wolf, ''Chrysocyon brachyurus'', in central Brazil". ''Journal of Zoology'', volume 240, issue 2, pp.  277–84. {{doi|10.1111/j.1469-7998.1996.tb05284.x}}</ref>

<ref name="moti1970">M. Motidome, M. E. Leekning, and O. R. Gottlieb (1970): "[http://memoria.bn.br/DocReader/DocReader.aspx?bib=158119&Pesq=sciencia&pagfis=15482 A química de Solanáceas brasileiras. 1 - A presenca de solamargina e de solasonina no juá e na lobeira.]" ''Anais da Academia Brasileira de Ciências'', volume 42, supplement, pp.  375–76</ref>

<ref name="lomb1993">J. A. Lombardi and J. C. Motta Jr. (1993): "[https://www.researchgate.net/profile/Jose-Motta-Junior-2/publication/283322828_Seed_dispersal_of_Solanum_lycocarpum_StHil_Solanaceae_by_the_Maned_Wolf_Chrysocyon_Brachyurus_Mammalia_Canidae/links/56335e0a08ae88cf81ba3a26/Seed-dispersal-of-Solanum-lycocarpum-StHil-Solanaceae-by-the-Maned-Wolf-Chrysocyon-Brachyurus-Mammalia-Canidae.pdf Seed dispersal of ''Solanum lycocarpum'' St. Hil. (Solanaceae) by the maned wolf, ''Chrysocyon brachyurus'' Illiger (Mammalia, Canidae).]" ''Ciência e Cultura'', volume 45, pp.  126–27.</ref>

<ref name=oliv1988>A.T. Oliveira-Filho and L. C. A. Oliveira (1988): "[https://www.bdpa.cnptia.embrapa.br/consulta/busca?id=552969 Biologia floral de uma população de ''Solanum lycocarpum'' St. Hil. (Solanaceae) em Lavras]". ''Revista Brasileira de Botânica, volume 11, pp.  23–32. Abstract accessed on 2020-06-30</ref>

<ref name=fari2010>Fabiane Farina (2010): "Glycemic and urinary volume responses in diabetic mellitus rats treated with ''Solanum Lycocarpum''." ''Applied Physiology, Nutrition, and Metabolism'', volume 35, issue 1, pp.  40–44. {{doi|10.1139/H09-131}}</ref>

<ref name="silv2015">Bicalho Silva and Viviane de Cassia (2015): "Larvicidal activity of oils, fatty acids, and methyl esters from ripe and unripe fruit of ''Solanum lycocarpum'' (Solanaceae) against the vector ''Culex quinquefasciatus'' (Diptera: Culicidae)". ''Rev. Soc. Bras. Med. Trop.'' '''48'''(5): 610-613 {{doi|10.1590/0037-8682-0049-2015}}{{open access}} {{PMID|26516974}}</ref>

<ref name=mira2012>Mariza A. Miranda, Lizandra G. Magalhaes, Renata F. J. Tiossi, Christian C. Kuehn, Luiz G. R. Oliveira, Vanderlei Rodrigues, James D. McChesney and Jairo K. Bastos (2012): "Evaluation of the schistosomicidal activity of the steroidal alkaloids from ''Solanum Lycocarpum'' fruits". ''Parasitology Research'', volume 111, pp.  257–62.  {{doi|10.1007/s00436-012-2827-8}}</ref>

<ref name=cler2011>M. T. P. S. Clerici, C. Kallmann, F. O. G. Gaspi, M. A. Morgano, F. Martinez-Bustos, and Y. K. Chang (2011): "Physical, chemical and technological characteristics of ''Solanum lycocarpum'' A. St. HILL (Solanaceae) fruit flour and starch". ''Food Research International'', volume 44, issue 7, pp.  2143–50. {{doi|10.1016/j.foodres.2011.01.060}}</ref>

<ref name=agno2000>Rodrigo Dall’Agnol and Gilsane L. von Poser (2000): "The use of complex polysaccharides in the management of metabolic diseases: the case of ''Solanum lycocarpum'' fruits". ''Journal of Ethnopharmacology'', volume 71, issues 1–2, pp.  337–41. {{doi|10.1016/S0378-8741(00)00165-3}}</ref>

<ref name=pasc2013>Aline M. Pascoal, Maria C. B. Di-Medeiros, Karla A. Batista, Maria I. G. Leles, Luciano Moraes Lião, and Kátia F. Fernandes (2013): "Extraction and chemical characterization of starch from ''S. lycocarpum'' fruits". ''Carbohydrate Polymers'', volume 98, issue 2, pp.  1304–10. {{doi|10.1016/j.carbpol.2013.08.009}}</ref>
  
<ref name=mora2015>Melissa G. Morais, Guilherme A. F. da Costa, Álan A. Aleixo, Graziela T. de Oliveira, Lucas F. Alves, and Joaquim M. Duarte-Almeida (2015): "Antioxidant, antibacterial and cytotoxic potential of the ripe fruits of ''Solanum lycocarpum'' A. St. Hil. (Solanaceae)". ''Journal of Natural Product Research'', volume 29, issue 5, pp.  480–83 {{doi|10.1080/14786419.2014.951930}}</ref>

<ref name=oliv2003>A. C. P. Oliveira, D. C. Endringer, R. J. P. Araújo, M. G. L. Brandão, and M. M. Coelho (2003): "The starch from ''Solanum lycocarpum'' St. Hill. fruit is not a hypoglycemic agent". ''Brazilian Journal of Medical and Biological Research'', volume 36, issue 4, pp.  525–30. {{doi|10.1590/S0100-879X2003000400017}}</ref>

<ref name=agui2005>Ludmilla M. de S. Aguiar (2005): "First record on the use of leaves of ''Solanum lycocarpum'' (Solanaceae) and fruits of ''Emmotum nitens'' (Icacinacea) by ''Platyrrhinus lineatus'' (E. Geoffroy) (Chiroptera, Phyllostomidae) in the Brazilian Cerrado". ''Revista Brasileira de Zoologia'', volume 22, issue 2, pp.  509–10.  {{doi|10.1590/S0101-81752005000200030}}</ref>

<ref name=jaeg1944>[[Edmund C. Jaeger]] (1944): ''[https://archive.org/details/in.ernet.dli.2015.350249/page/n153/mode/1up A Source-Book Of Biological Names And Terms]''. pp.  41, 153.  Published by Charles C. Thomas, Illinois.</ref>

<ref name=mart2015>Gilmarcio Z. Martins, Raquel R. D. Moreira, Cleopatra S. Planeta, Adélia E. Almeida, Jairo K. Bastos, Lígia Salgueiro, Carlos Cavaleiro, and Maria do Céu Sousa  (2015): "Effects of the extract and glycoalkaloids of ''Solanum lycocarpum'' St. Hill on ''Giardia lamblia'' trophozoites" ''Pharmacognosy Magazine'', volume 11, supplement 1, pp.  S161–65. {{doi|10.4103/0973-1296.157721|doi-access=free}} {{PMID|26109762}}</ref>

<ref name=pere2019>Ana P. A. Pereira, Célio F. F. Angolini, Bruno N. Paulino, Leonardo B. C. Lauretti, Eduardo A. Orlando, Joyce G. S. Silva, Iramaia A. Neri-Numa, Jane D. R. P. Souza, Juliana A. L. Pallone, Marcos N. Eberlin, and Glaucia M. Pastore (2019): "A comprehensive characterization of Solanum lycocarpum St. Hill and Solanum oocarpum Sendtn: Chemical composition and antioxidant properties". ''Food Research International'', volume 124, pp.  61–69. {{doi|10.1016/j.foodres.2018.09.054}}</ref>

<ref name=mira2013>Mariza A. Miranda,  Renata F. J. Tiossi,  Mariana R. da Silva,  Kelly C. Rodrigues,  Christian C. Kuehn,  Luiz G. R. Oliveira,  Sérgio Albuquerque,   James D. McChesney,  Claudio M. Lezama‐Davila,  Angelica P. Isaac‐Marquez, and  Jairo K. Bastos (2013): "In vitro Leishmanicidal and cytotoxic activities of the glycoalkaloids from ''Solanum lycocarpum'' (Solanaceae) fruits". ''Chemistry & Biodiversity'', volume 10, issue 4, pp.  642–48.{{doi|10.1002/cbdv.201200063}}</ref>

<ref name=pint2007>Lilian V. A. Pinto, Edvaldo A. A. Da Silva, Antonio C. Davide, Valquíria A. Mendes De Jesus, Peter E. Toorop, Henk W. M. Hilhorst (2007): "Mechanism and control of ''Solanum lycocarpum'' seed germination". ''Annals of Botany'', volume 100, issue 6, pp.  1175–1187. {{doi|10.1093/aob/mcm211}}</ref>

<ref name=clem2018>L. C. Clementino,  A. M. A. Velásquez, T. G. Passalacqua, L. Almeida, M. A. S. Graminha, G. Z. Martins, L. Salgueiro, C. Cavaleiro, M. C. Sousa, R. R. D. Moreira (2018): "In vitro activities of glycoalkaloids from the ''Solanum lycocarpum'' against ''Leishmania infantum''". ''Revista Brasileira de Farmacognosia'', volume 28, issue 6, pp.  673–77. {{doi|10.1016/j.bjp.2018.07.008}}</ref>

<ref name=soar2019>M. A. Soares, L. P. Sánchez, J. B. Isaac Júnior, E. S. Pereira, E. M.s Pires, G. L. D. Leite, and J. C. Zanuncio (2019): "''Solanum lycocarpum'' Saint Hilaire (Solanaceae) is host plant of ''Leucanella memusae'' (Walker) (Lepidoptera: Saturniidae: Hemileucinae) in Brazilian mountain grasslands". ''Florida Entomologist'', volume 102, issue 3, pp.  624–26. {{doi|10.1653/024.102.0341}}</ref>

<ref name=souz1998>A. L. T. De Souza, G. W. Fernandes, J. E. C. Figueira, and M. O. Tanaka (1998): "Natural history of a gall-inducing weevil ''Collabismus clitellae'' (Coleoptera: Curculionidae) and some effects on its host plant ''Solanum lycocarpum'' (Solanaceae) in Southeastern Brazil". ''Annals of the Entomological Society of America'', volume 91, issue 4, pp.  404–09. {{doi|10.1093/aesa/91.4.404}}</ref>

</references>

{{commons category|Solanum lycocarpum}}

{{Taxonbar|from=Q3280462}}

[[Category:Solanum|lycocarpum]]
[[Category:Flora of Brazil]]
[[Category:Flora of the Cerrado]]
[[Category:Tropical fruit]]
[[Category:Edible Solanaceae]]