Solanum lycocarpum

Solanum lycocarpum, or wolf apple, is common in the Brazilian savanna, the Cerrado ecoregion. The plant is called lobeira ("wolf's plant") or fruta-do-lobo ("wolf's fruit") in Portuguese.[1]

Not to be confused with Solanum lycopersicum, wolfberry, or Wolf River (apple).

Solanum lycocarpum
Scientific classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Solanales
Family: Solanaceae
Genus: Solanum
Species:
S. lycocarpum
Binomial name
Solanum lycocarpum

The name "wolf apple" comes from the fact that they account for more than 50% of the Maned wolf's diet.[1] Likewise, the scientific name "lycocarpum" is formed from Latinized Greek elements "lyco-", meaning "wolf", and "carpum" meaning "fruit".[2]

Range

The native range of the wolf apple tree is the Brazilian savannah, but it grows also on pastures and disturbed land, such as highway margins, in various parts of Brazil.[3] It prefers moist, clay soil, full sun, and mild temperatures.[4][5]

Description
Flower
Fruit

The wolf apple plant is a flowering shrub or small tree with round open crown, ranging in height from 1.2 to 5 m. The leaves are large (16 to 28 cm 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 sepals, five blue or purple petals fused at the base, and five large yellow anthers that release pollen through an opening at the tip. The flowers are arranged in helical 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 bees, including Xylocopa virescens, X. frontalis, Oxaea flavescens, Centris scopipes, C. fuscata, Bombus morio, Exomalopsis sp., Pseudaugochloropsis graminea, Apis mellifera, and Megachile benigna.[4][5]

Fruit production occurs mostly between December and January. The fruits are large, up to 20 cm in diameter and weighing up to over 900 g, and contain up to 500 dark brown comma-shaped seeds, about 7 × 5 × 2 mm.[6] The rind is thin and lustrous, and remains green even after ripening.[6] The pulp is yellowish, soft, sweetish and extremely aromatic.[3] They resemble the tomato in external shape and internal morphology,[5] but also the eggplant for the texture and color of the flesh. The maned wolf is an important seed-spreading agent.[7]

Uses

Food

The ripe fruits are edible to humans, and are consumed by local populations in jams and preserves, but the unripe fruits are quite rich in tannin.[5] 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.[3] The dry pulp, without peel and seeds, yields over 50% starch, more than cassava.[8] The starch is locally separated by blending the fruit with water and straining.[3]

The pulp of the ripe fruit contains about 75% moisture, 11% carbohydrate, 1% fat, 1% protein, 1% ash, 2.5% 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.[9]

All other parts of the plant are poisonous to humans. However, the leaves are eaten by cattle, and are also an important food source for the bat Platyrrhinus lineatus.[10]

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.[5] About 30% of the starch extracted from almost-ripe fruits, which commercialized as capsules, is digestion-resistant,[3] which seems to be the rationale for these popular uses.[11] High-fiber Solanum lycocarpum flour was found to reduce diabetes symptoms in diabetic rats;[12] however, the starch did not show such effect.[13]

The fruit contains about 1.5% of glycoalkaloids in its dried fruits, chiefly solamargine and solasonine,[14] mostly in the peel and seeds.[3] Extracts have been shown to kill in vitro the human parasites Schistosoma mansoni[15] and Giardia lamblia,[16] Leishmania amazonensis,[17] and Leishmania infantum.[18] The main phenolic compounds in ethanol extracts of the ripe fruits are caffeic acid and chlorogenic acid.[19]

It is believed that the fruits protect the maned wolf from infection of the giant kidney worm, which is usually fatal for the animal.[5]

Solanum lycocarpum fruit extracts have also been shown to kill larvae of the Culex mosquito.[20]

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.[21]

The weevil Collabismus clitellae lays eggs into stems of the plant, causing the formation of galls, with measurable harmful effect on the plant's growth and fruit production.[22]

References
  1. 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, pages 277–284. doi:10.1111/j.1469-7998.1996.tb05284.x
  2. Edmund C. Jaeger (1944): A Source-Book Of Biological Names And Terms. Pages 41 and 153. Published by Charles C. Thomas, Illinois.
  3. 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, pages 2143-2150. doi:10.1016/j.foodres.2011.01.060
  4. A.T. Oliveira-Filho and L. C. A. Oliveira (1988): "Biologia floral de uma população de Solanum lycocarpum St. Hil. (Solanaceae) em Lavras". Revista Brasileira de Botânica, volume 11, pages 23–32. Abstract accessed on 2020-06-30
  5. Fernando Tatagiba (2006): "Lobeira, fruta-de-lobo". Online article on the Biólogo website, published 2006-10-06. Accessed on 2020-06-30.
  6. 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, pages 1175–1187. doi:10.1093/aob/mcm211
  7. J. A. Lombardi and J. C. Motta Jr. (1993): "Seed dispersal of Solanum lycocarpum St. Hil. (Solanaceae) by the maned wolf, Chrysocyon brachyurus Illiger (Mammalia, Canidae)." Ciência e Cultura, volume 45, pages 126–127.
  8. 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, pages 1304-1310. doi:10.1016/j.carbpol.2013.08.009
  9. 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, pages 61-69. doi:10.1016/j.foodres.2018.09.054
  10. 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, pages 509-510. doi:10.1590/S0101-81752005000200030
  11. 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, pages 337-341. doi:10.1016/S0378-8741(00)00165-3
  12. 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, pages 40-44. doi:10.1139/H09-131
  13. 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, pages 525-530. doi:10.1590/S0100-879X2003000400017
  14. M. Motidome, M. E. Leekning, and O. R. Gottlieb (1970): "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, pages 375–376
  15. 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, pages 257–262. doi:10.1007/s00436-012-2827-8
  16. 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, suplement 1, pages S161–S165. doi:10.4103/0973-1296.157721 PMID 26109762
  17. 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, pages 642-648.doi:10.1002/cbdv.201200063
  18. 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, pages 673-677. doi:10.1016/j.bjp.2018.07.008
  19. 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, pages 480-483 doi:10.1080/14786419.2014.951930
  20. 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)".
  21. 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, pages 624-626. doi:10.1653/024.102.0341
  22. 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, pages 404–409. doi:10.1093/aesa/91.4.404
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