Solanum lycocarpum, or wolf apple, is common in the Braziliansavanna, the Cerradoecoregion. The plant is called lobeira ("wolf's plant") or fruta-do-lobo ("wolf's fruit") in Portuguese.[1]
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 LatinizedGreek elements "lyco-", meaning "wolf", and "carpum" meaning "fruit".[2]
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]
FlowerFruit
The wolf apple plant is a flowering shrub or small tree with round open crown, ranging in height from 1.2 to 5m (4 to 16ft). The large leaves are 16 to 28cm (6 to 11in) 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 20cm (8in) in diameter and weighing 900g (2lb) or more, and contain up to 500 dark brown comma-shaped seeds, about 7mm ×5mm ×2mm (1⁄4in ×3⁄16in ×1⁄16in).[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]
Culinary use
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]
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 wolf apple tree is the natural host of the larva of the butterflyLeucanella memusae. Contact with the urticating bristles of this caterpillar may result in severe dermatitis and hemorrhage.[21]
The weevilCollabismus 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]
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
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
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
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
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
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
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
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
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
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:10.4103/0973-1296.157721PMID26109762
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
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
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
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:10.1590/0037-8682-0049-2015PMID26516974
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
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