''Solanum lycocarpum'', or wolf apple, is common in the

Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...

ian

savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to ...

, the

Cerrado The ''Cerrado'' (, ) is a vast ecoregion of tropical savanna in eastern Brazil, particularly in the states of Goiás, Mato Grosso do Sul, Mato Grosso, Tocantins, Minas Gerais, and the Federal District. The core areas of the Cerrado biome are t ...

ecoregion An ecoregion (ecological region) or ecozone (ecological zone) is an ecologically and geographically defined area that is smaller than a bioregion, which in turn is smaller than a biogeographic realm. Ecoregions cover relatively large areas of l ...

. The plant is called ''lobeira'' ("wolf's plant") or ''fruta-do-lobo'' ("wolf's fruit") in

Portuguese Portuguese may refer to: * anything of, from, or related to the country and nation of Portugal ** Portuguese cuisine, traditional foods ** Portuguese language, a Romance language *** Portuguese dialects, variants of the Portuguese language ** Portu ...

. The name "wolf apple" comes from the fact that they account for more than 50% of the

maned wolf The maned wolf (''Chrysocyon brachyurus'') is a large canine of South America. It is found in Argentina, Brazil, Bolivia, Peru, and Paraguay, and is almost extinct in Uruguay. Its markings resemble those of foxes, but it is neither a fox nor a w ...

's diet. Likewise, the scientific name "''lycocarpum''" is formed from

Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...

ized

Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...

elements "''lyco-''", meaning "wolf", and "''carpum''" meaning "fruit".

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. It prefers moist, clay soil, full sun, and mild temperatures.

Description

Flickr - João de Deus Medeiros - Solanum lycocarpum (1)

The wolf apple plant is a flowering

shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...

or small tree with round open crown, ranging in height from . The large leaves are 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 A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coined b ...

s, five blue or purple

petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...

s fused at the base, and five large yellow

anther The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...

s that release

pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...

through an opening at the tip. The flowers are arranged in helical monochasal cyme, opening from the base up. Flowers lack

nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...

, and are either male or hermaphroditic. The latter produce fruit after cross-pollination by several species of pollen-collecting

bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...

s, including '' Xylocopa virescens'', '' X. frontalis'', '' Oxaea flavescens'', ''

Centris scopipes The genus ''Centris'' contains circa 250 species of large apid bees occurring in the Neotropical and Nearctic realms, from Kansas to Argentina. Most females of these bees possess adaptations for carrying floral oils rather than (or in addition ...

'', '' C. fuscata'', ''

Bombus morio ''Bombus morio'' is one of the few bumblebee species found in South America. These bees reside mainly in the forests of Brazil, nesting on the surface of the ground. They are one of the biggest species of bumblebee and are important pollinators. ...

'', ''

Exomalopsis ''Exomalopsis'' is a genus of bees in the family Apidae. They occur in the Western Hemisphere (Neotropical and Nearctic realms).Velez-Ruiz, R. I. and A. H. Smith-Pardo. (2013)New species of ''Exomalopsis'' and its associated cleptoparasite ''No ...

'' sp., '' Pseudaugochloropsis graminea'', ''

Apis mellifera The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...

'', and ''

Megachile benigna ''Megachile benigna'' is a species of bee in the family Megachilidae Megachilidae is a cosmopolitan family of mostly solitary bees. Both that their pollen-carrying structure (called a '' scopa'') is restricted to the ventral surface of the ...

''. Fruit production occurs mostly between December and January. The fruits are large, up to in diameter and weighing or more, and contain up to 500 dark brown comma-shaped seeds, about . The rind is thin and lustrous, and remains green even after ripening. The pulp is yellowish, soft, sweetish and extremely aromatic. They resemble the

tomato The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...

in external shape and internal morphology, but also the

eggplant Eggplant ( US, Canada), aubergine ( UK, Ireland) or brinjal (Indian subcontinent, Singapore, Malaysia, South Africa) is a plant species in the nightshade family Solanaceae. ''Solanum melongena'' is grown worldwide for its edible fruit. Mos ...

for the texture and color of the flesh. The maned wolf is an important seed-spreading agent.

Uses

Food

The ripe fruits are edible to

humans Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...

, and are consumed by local populations in jams and preserves, but the unripe fruits are quite rich in

tannin Tannins (or tannoids) are a class of astringent, polyphenolic biomolecules that bind to and precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' (from Anglo-Norman ''tanner'', ...

. The fruits are used as a fallback source of food for

cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult mal ...

in the dry season. At the onset of maturation, the whole fruit contains 85% moisture (by weight), 5% starch, and 10% other residue. The dry pulp, without peel and seeds, yields over 50% starch, more than

cassava ''Manihot esculenta'', common name, commonly called cassava (), manioc, or yuca (among numerous regional names), is a woody shrub of the spurge family, Euphorbiaceae, native to South America. Although a perennial plant, cassava is extensively ...

. The starch is locally separated by blending the fruit with water and straining. The pulp of the ripe fruit contains about 75% moisture, 11%

carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or ma ...

, 1%

fat In nutrition science, nutrition, biology, and chemistry, fat usually means any ester of fatty acids, or a mixture of such chemical compound, compounds, most commonly those that occur in living beings or in food. The term often refers spec ...

, 1%

protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...

, 1% ash, 2.5%

soluble fiber Dietary fiber (in British English fibre) or roughage is the portion of plant-derived food that cannot be completely broken down by human digestive enzymes. Dietary fibers are diverse in chemical composition, and can be grouped generally by the ...

, and 2% insoluble fiber. The main volatiles in the ripe fruit are

hexanal Hexanal, also called hexanaldehyde or caproaldehyde is an alkyl aldehyde used in the flavor industry to produce fruity flavors. Its scent resembles freshly cut grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monoc ...

(which has the scent of grass) and ethyl butanoate (scent of apple), although dozens of other compounds contribute to the fruit's aroma. All other parts of the plant are

poison Poison is a chemical substance that has a detrimental effect to life. The term is used in a wide range of scientific fields and industries, where it is often specifically defined. It may also be applied colloquially or figuratively, with a broa ...

ous to humans. However, the leaves are eaten by

, and are also an important food source for the

bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...

'' Platyrrhinus lineatus''.

Medicine

Currently, there are no proven medicinal uses for the wolf apple, but the plant is used in the local

folk medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...

for

obesity Obesity is a medical condition, sometimes considered a disease, in which excess body fat has accumulated to such an extent that it may negatively affect health. People are classified as obese when their body mass index (BMI)—a person's we ...

, cholesterol reduction, and as a treatment for

diabetes Diabetes, also known as diabetes mellitus, is a group of metabolic disorders characterized by a high blood sugar level ( hyperglycemia) over a prolonged period of time. Symptoms often include frequent urination, increased thirst and increased ap ...

. About 30% of the starch extracted from almost-ripe fruits, which commercialized as capsules, is digestion-resistant, which seems to be the rationale for these popular uses. High-fiber ''Solanum lycocarpum'' flour was found to reduce diabetes symptoms in diabetic rats; however, the starch did not show such effect. The fruit contains about 1.5% of

glycoalkaloid Glycoalkaloids are a family of chemical compounds derived from alkaloids to which sugar groups are appended. Several are potentially toxic, most notably the poisons commonly found in the plant species ''Solanum dulcamara'' (bittersweet nightshade) ...

s in its dried fruits, chiefly

solamargine Solamargine is a cytotoxic chemical compound that occurs in plants of the family Solanaceae, such as potatoes, tomatoes, and eggplants. It has been also isolated from '' Solanum nigrum'' fungal endophyte ''Aspergillus flavus''. It is a glycoalk ...

and

solasonine Solasonine is a glycoalkaloid that is found in Solanum plants of the family Solanaceae. Solasonine is a poisonous chemical compound when used at high levels. It is a glycoside of solasodine Solasodine is a poisonous alkaloid chemical compound ...

, mostly in the peel and seeds. Extracts have been shown to kill

in vitro ''In vitro'' (meaning in glass, or ''in the glass'') studies are performed with microorganisms, cells, or biological molecules outside their normal biological context. Colloquially called "test-tube experiments", these studies in biology an ...

the human parasites ''

Schistosoma mansoni A paired couple of ''Schistosoma mansoni''. ''Schistosoma mansoni'' is a water-borne parasite of humans, and belongs to the group of blood flukes (''Schistosoma''). The adult lives in the blood vessels ( mesenteric veins) near the human inte ...

'' and ''

Giardia lamblia ''Giardia duodenalis'', also known as ''Giardia intestinalis'' and ''Giardia lamblia'', is a flagellated parasitic microorganism of the genus '' Giardia'' that colonizes the small intestine, causing a diarrheal condition known as giardiasis. Th ...

'', ''

Leishmania amazonensis ''Leishmania amazonensis'' is a parasite responsible for the disease leishmaniasis. This species has been known to spread by using sandflies Sandfly (or sand fly) is a colloquial name for any species or genus of flying, biting, blood-suckin ...

'', and ''

Leishmania infantum ''Leishmania infantum'' is the causative agent of infantile visceral leishmaniasis in the Mediterranean region and in Latin America, where it has been called ''Leishmania chagasi''. It is also an unusual cause of cutaneous leishmaniasis, which i ...

''. The main phenolic compounds in

ethanol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an Alcohol (chemistry), alcohol with the chemical formula . Its formula can be also written as or (an ethyl ...

extracts of the ripe fruits are

caffeic acid Caffeic acid is an organic compound that is classified as a hydroxycinnamic acid. This yellow solid consists of both phenolic and acrylic functional groups. It is found in all plants because it is an intermediate in the biosynthesis of lignin, one ...

and

chlorogenic acid Chlorogenic acid (CGA) is the ester of caffeic acid and (−)-quinic acid, functioning as an intermediate in lignin biosynthesis. The term "chlorogenic acids" refers to a related polyphenol family of esters, including hydroxycinnamic acids (caffe ...

. It is believed that the fruits protect the

from infection of the giant kidney worm, which is usually fatal for the animal. ''Solanum lycocarpum'' fruit extracts have also been shown to kill larvae of the ''Culex''

mosquito Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small flies within the family Culicidae (from the Latin ''culex'' meaning " gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish for "li ...

Pests

The wolf apple tree is the natural host of the larva of the

butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...

Leucanella memusae ''Leucanella'' is a genus of moths in the family Saturniidae first described by Claude Lemaire in 1969. Species The genus includes the following species: *'' Leucanella acutissima'' (Walker, 1865) *'' Leucanella anikae'' Meister & Brechlin, 20 ...

''. Contact with the urticating bristles of this caterpillar may result in severe

dermatitis Dermatitis is inflammation of the skin, typically characterized by itchiness, redness and a rash. In cases of short duration, there may be small blisters, while in long-term cases the skin may become thickened. The area of skin involved can v ...

and

hemorrhage Bleeding, hemorrhage, haemorrhage or blood loss, is blood escaping from the circulatory system from damaged blood vessels. Bleeding can occur internally, or externally either through a natural opening such as the mouth, nose, ear, urethra, vag ...

. The

weevil Weevils are beetles belonging to the Taxonomic rank, superfamily Curculionoidea, known for their elongated snouts. They are usually small, less than in length, and Herbivore, herbivorous. Approximately 97,000 species of weevils are known. They b ...

'' Collabismus clitellae'' lays eggs into stems of the plant, causing the formation of

gall Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants, fungi, or animals. Plant galls are abnormal outgrowths of plant tissues, similar to be ...

s, with measurable harmful effect on the plant's growth and fruit production.

References

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Lobeira, fruta-de-lobo
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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 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. 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)". 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. 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. 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. 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. 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 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. 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.

Edmund C. Jaeger Edmund Carroll Jaeger, D.Sc.,University of California, Riverside, Science Library (January 28, 1887 – August 2, 1983) was an American biologist known for his works on desert ecology. He was born in Loup City, Nebraska to Katherine (née Gunther ...

(1944):
A Source-Book Of Biological Names And Terms
'. pp. 41, 153. Published by Charles C. Thomas, Illinois. 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, pp. S161–65. 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. 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. 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. 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. 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. 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. {{Taxonbar, from=Q3280462 lycocarpum Flora of Brazil Flora of the Cerrado Tropical fruit Edible Solanaceae