Inga edulis
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''Inga edulis'', known as ice-cream bean, ice-cream-bean, joaquiniquil, cuaniquil, guama or guaba, is a fruit native to
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
. It is in the mimosoid tribe of the legume family
Fabaceae The Fabaceae or Leguminosae,International Code of Nomenc ...
. It is widely grown, especially by
Indigenous Indigenous may refer to: *Indigenous peoples *Indigenous (ecology), presence in a region as the result of only natural processes, with no human intervention *Indigenous (band), an American blues-rock band *Indigenous (horse), a Hong Kong racehorse ...
Amazonians, for shade, food, timber, medicine, and production of the alcoholic beverage cachiri. It is popular in
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = Seal (emblem), National seal , national_motto = "Fi ...
,
Ecuador Ecuador ( ; ; Quechua: ''Ikwayur''; Shuar: ''Ecuador'' or ''Ekuatur''), officially the Republic of Ecuador ( es, República del Ecuador, which literally translates as "Republic of the Equator"; Quechua: ''Ikwadur Ripuwlika''; Shuar: ''Eku ...
,
Pernambuco Pernambuco () is a state of Brazil, located in the Northeast region of the country. With an estimated population of 9.6 million people as of 2020, making it seventh-most populous state of Brazil and with around 98,148 km², being the 19 ...
-
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 ...
,
Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...
and
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the Car ...
. The taxonomic name ''Inga'' is derived from its name with the
Tupí people A subdivision of the Tupi-Guarani linguistic families, the Tupi people were one of the largest groups of indigenous Brazilians before its colonization. Scholars believe that while they first settled in the Amazon rainforest, from about 2,900 ...
of South America, while the species name '' edulis'' is Latin for "edible". The common name "ice-cream bean" alludes to the sweet flavor and smooth texture of the pulp.


Biology

Mature trees of ''Inga edulis'' reach 30 m (98 ft) high and 60 cm (2.0 ft) diameter at breast height, usually branching from below 3 m (9.8 ft). The branches form a broad, flat, moderately dense canopy. ''Inga edulis'' can be evergreen in tropical regions or deciduous when planted in colder regions. The tree has a pale grey coloured trunk. The stems and young twigs can be sparsely to densely haired. The leaves are alternate, evenly
pinnate Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and in ...
, 10–30 cm long with 4–6 pairs of opposite, dark-green, membranous, slightly pubescent, oval leaflets. The terminal leaflets can grow up to 18 cm long by 11 cm wide in comparison to the basal ones. Extrafloral
nectaries 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 ...
are placed on petioles and
stipule In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole). Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many speci ...
s can be either inconspicuous, absent, or caduceus. Inga species are in symbiosis with ants (e.g.,
Pheidole ''Pheidole'' is a genus of ants that belongs to the ant subfamily Myrmicinae. The genus is widespread and ecologically dominant. It probably includes more than 1000 species. The genus first evolved in the Americas, eventually spreading across th ...
s spp.), which get the nectar of the extrafloral nectaries. In exchange the ants will patrol over the Inga plant to protect it against
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
s. There are certain insects such as Riodinid caterpillars which excrete sugary honeydew from their tentacle nectaries. The ants will form a symbiosis with the caterpillars by letting them feed on the tree and in favour of getting sugary honeydew from them. Flowers are fragrant, sessile, pentamerous and are arranged in dense axillary spikes. The flower has a calyx tube with five puberulent, striate lobes, corolla with five silky, villous petals. These are around 14–20 mm long and contain numerous white
stamen 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 filame ...
s. The fruits are longitudinally ribbed, cylindrical indehiscent
leguminous A legume () is a plant in the family Fabaceae (or Leguminosae), or the fruit or seed of such a plant. When used as a dry grain, the seed is also called a pulse. Legumes are grown agriculturally, primarily for human consumption, for livestock for ...
pods which can be straight, curved or often spirally twisted up. They are pendant and up to 1 m long and yellowish brown to greenly coloured. The amount of ovoid seeds can vary from 10 to 20, which are purplish-black to olive colored. These are embedded in the sweet, cottony, white
arillus An aril (pronounced ), also called an arillus, is a specialized outgrowth from a seed that partly or completely covers the seed. An arillode or false aril is sometimes distinguished: whereas an aril grows from the attachment point of the see ...
which gives it the name Icecream bean, since they taste sweet. ;Symbiosis: The Inga plant can also form symbiotic relationships where nitrogen gas can be fixated by
rhizobia Rhizobia are diazotrophic bacteria that fix nitrogen after becoming established inside the root nodules of legumes (Fabaceae). To express genes for nitrogen fixation, rhizobia require a plant host; they cannot independently fix nitrogen. In gene ...
l bacteria and
mycorrhiza   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...
as other legumes can. Surveys have shown that ''Inga edulis'' undergo a mutualistic relationships with the bacterial strain of ''
Bradyrhizobium ''Bradyrhizobium'' is a genus of Gram-negative soil bacteria, many of which fix nitrogen. Nitrogen fixation is an important part of the nitrogen cycle. Plants cannot use atmospheric nitrogen (N2); they must use nitrogen compounds such as nitrat ...
''. File:Inga Flowers.JPG, Flowers File:Inga edulis, mature and juvenile Guama pods (10108341614).jpg, Mature and young pods


Synonyms

This plant has a convoluted history of
synonymy A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
with ''
Inga vera ''Inga vera'' is a species of tropical tree in the family Fabaceae The Fabaceae or Leguminosae,Brenan Brenan is a surname and a given name. Notable people with the name include: Surname * Gerald Brenan (1894 – 1987), British writer and Hispanist * James Brenan (1837 – 1907), Irish artist * Ryan Brenan (1798 – 1868), Australian politician fr ...
and
Kunth Carl Sigismund Kunth (18 June 1788 – 22 March 1850), also Karl Sigismund Kunth or anglicized as Charles Sigismund Kunth, was a German botanist. He is known for being one of the first to study and categorise plants from the American continents, ...
are actually ''I. edulis'', whereas that based on the writings of
Carl Ludwig Willdenow Carl Ludwig Willdenow (22 August 1765 – 10 July 1812) was a German botanist, pharmacist, and plant taxonomist. He is considered one of the founders of phytogeography, the study of the geographic distribution of plants. Willdenow was als ...
refers to the actual ''I. vera''. ''Inga edulis'' in works referring back to authorities other than
von Martius Carl Friedrich Philipp (Karl Friedrich Philipp) von Martius (17 April 1794 – 13 December 1868) was a German botanist and explorer. Life Martius was born at Erlangen, the son of Prof Ernst Wilhelm Martius, court apothecary. He graduated PhD ...
usually refers to ''
Inga feuilleei ''Inga feuilleei'' (named after Louis Feuillée), commonly known as pacay or ice-cream bean tree, is a tree in the family Fabaceae native to Andes, Andean valleys of northwestern South America. Pacay trees, as is the case with other trees in g ...
''.ILDIS (2005) Synonyms of ''Inga edulis'' Mart.: *''Feuilleea edulis'' (Mart.) Kuntze *''Inga benthamiana''
Meisn. Carl Daniel Friedrich Meissner (1 November 1800 – 2 May 1874) was a Swiss botanist. Biography Born in Bern, Switzerland on 1 November 1800, he was christened Meisner but later changed the spelling of his name to Meissner. For most of his 40 ...
*''Inga edulis''
var. In botanical nomenclature, variety (abbreviated var.; in la, varietas) is a taxonomic rank below that of species and subspecies, but above that of form. As such, it gets a three-part infraspecific name. It is sometimes recommended that the ...
''grenadensis'' Urb. *''Inga minutula'' (Schery) T.S.Elias *''Inga scabriuscula'' Benth. *''Inga vera'' Kunth *''Inga vera'' ''sensu'' Brenan *''Inga ynga'' (Vell.) J.W.Moore *''Mimosa inga'' L. *''Mimosa ynga'' Vell.


Ecology

The natural distribution of ''Inga edulis'' spreads from
Central Central is an adjective usually referring to being in the center of some place or (mathematical) object. Central may also refer to: Directions and generalised locations * Central Africa, a region in the centre of Africa continent, also known as ...
to
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
and ranges from subtropical dry to tropical wet conditions. It can be found at elevations from sea level up to 2200 m. ''Inga edulis'' grows best when mean annual daytime temperature ranges from 23 to 30 °C. However, it can tolerate 18–35 °C. During the resting period mature trees can survive −2 °C while young plants are killed at 0 °C. Preferred
annual precipitation In meteorology, precipitation is any product of the condensation of atmospheric water vapor that falls under gravitational pull from clouds. The main forms of precipitation include drizzle, rain, sleet, snow, ice pellets, graupel and hai ...
ranges from 1200 to 2500 mm but 640 to 4000 mm of annual precipitation can be tolerated also. ''Inga edulis'' can be grown on a widely varying range of soil conditions. It prefers a
soil pH Soil pH is a measure of the acidity or basicity (alkalinity) of a soil. Soil pH is a key characteristic that can be used to make informative analysis both qualitative and quantitatively regarding soil characteristics. pH is defined as the neg ...
of 5–6.5 but can also grow in very strongly acidic soil down to a pH of 4.5 or moderately alkaline conditions up to pH 8. The natural habitat of ''Inga edulis'' includes margins of large rivers like the
Amazon Amazon most often refers to: * Amazons, a tribe of female warriors in Greek mythology * Amazon rainforest, a rainforest covering most of the Amazon basin * Amazon River, in South America * Amazon (company), an American multinational technology c ...
, thickets below high water line and wooded swamps. Therefore, the tree tolerates temporarily waterlogged soils for 2–3 months each year. However, it can also tolerate drought seasons up to 6 months. Further ''Inga edulis'' can cope with heavy soils or slope fairly well. In terms of soil fertility, ''Inga edulis'' is not only unpretentious but has the potential to improve soil quality. As a leguminous tree, ''Inga edulis'' can compensate for nutrient depleted soils by
fixing nitrogen Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. Atmos ...
with its root nodules. Ammonium and nitrates found in soil typically result in nodulation reduction. ''Inga edulis i''s an exception, as its nodules increase in size when ammonium levels are at 5mM. Nitrate on the other hand did not benefit ''Inga edulis'' and continued to show properties that prevented nodulation. Additionally to nitrogen, ''Inga edulis'' is reported to significantly increase the extractable
Phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...
content in the soil and thus further ameliorates degraded soils.


Cultivation

''Inga edulis'' is widely cultivated in
agroforestry Agroforestry is a land use management system in which trees or shrubs are grown around or among crops or pastureland. Trees produce a wide range of useful and marketable products from fruits/nuts, medicines, wood products, etc. This intentional ...
systems in its
neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...
dispersal area. This form of cultivation, often associated with
coffee Coffee is a drink prepared from roasted coffee beans. Darkly colored, bitter, and slightly acidic, coffee has a stimulant, stimulating effect on humans, primarily due to its caffeine content. It is the most popular hot drink in the world. S ...
or
cocoa Cocoa may refer to: Chocolate * Chocolate * ''Theobroma cacao'', the cocoa tree * Cocoa bean, seed of ''Theobroma cacao'' * Chocolate liquor, or cocoa liquor, pure, liquid chocolate extracted from the cocoa bean, including both cocoa butter and ...
culture, is widely known since
pre-Columbian In the history of the Americas, the pre-Columbian era spans from the original settlement of North and South America in the Upper Paleolithic period through European colonization, which began with Christopher Columbus's voyage of 1492. Usually, th ...
times.
Germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fer ...
of ''Inga edulis'' seeds is fairly easy as the seeds are recalcitrant and often germinate already in the pod. However, seeds lose their ability to germinate after two weeks of storage. Direct sowing of the seeds is possible, but the establishment of seedlings with already a certain height can decrease
competition Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indivi ...
with noxious weeds and thus increase growth. Further, the inoculation with
rhizobia Rhizobia are diazotrophic bacteria that fix nitrogen after becoming established inside the root nodules of legumes (Fabaceae). To express genes for nitrogen fixation, rhizobia require a plant host; they cannot independently fix nitrogen. In gene ...
and
mycorrhiza   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...
l fungi in depleted soils is recommended to promote growth. These inoculi can easily be produced by collecting soil, nodules and fine roots from mature, nodulated inga edulis stands. ''Inga edulis'' seems to be very resistant to pests and diseases. Minor damages can result of fungal attacks in the seedling stage. Mature ''Inga edulis'' can be defoliated by
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
larvae. Also, fruit fly larvae often damage the seed testa. In
Ecuador Ecuador ( ; ; Quechua: ''Ikwayur''; Shuar: ''Ecuador'' or ''Ekuatur''), officially the Republic of Ecuador ( es, República del Ecuador, which literally translates as "Republic of the Equator"; Quechua: ''Ikwadur Ripuwlika''; Shuar: ''Eku ...
, Inga edulis in specially susceptible to
mistletoe Mistletoe is the common name for obligate hemiparasitic plants in the order Santalales. They are attached to their host tree or shrub by a structure called the haustorium, through which they extract water and nutrients from the host plant. ...
infestations.


Uses

Food crop ''Inga edulis'' has been cultivated as a
fruit tree A fruit tree is a tree which bears fruit that is consumed or used by animals and humans — all trees that are flowering plants produce fruit, which are the ripened ovaries of flowers containing one or more seeds. In horticultural usage, ...
for millennia and is widely sold on the local South American marketplace, mainly for the sweet, succulent pulp surrounding the seeds. The white pulp (
aril An aril (pronounced ), also called an arillus, is a specialized outgrowth from a seed that partly or completely covers the seed. An arillode or false aril is sometimes distinguished: whereas an aril grows from the attachment point of the see ...
) is consumed raw as a sweet snack, though it is less nutritious than the seeds. Toxic compounds such as
trypsin Trypsin is an enzyme in the first section of the small intestine that starts the digestion of protein molecules by cutting these long chains of amino acids into smaller pieces. It is a serine protease from the PA clan superfamily, found in the dig ...
and
chymotrypsin Chymotrypsin (, chymotrypsins A and B, alpha-chymar ophth, avazyme, chymar, chymotest, enzeon, quimar, quimotrase, alpha-chymar, alpha-chymotrypsin A, alpha-chymotrypsin) is a digestive enzyme component of pancreatic juice acting in the duodenu ...
inhibitors contained in the seeds of ''Inga edulis'' are destroyed through cooking.The taste is described as resembling that of
vanilla ice cream Vanilla is frequently used to flavor ice cream, especially in North America, Asia, and Europe. Vanilla ice cream, like other flavors of ice cream, was originally created by cooling a mixture made of cream, sugar, and vanilla above a container of ...
. Some varieties even possess a slight cinnamon flavor. The seeds are only edible when cooked and have a taste similar to that of
chickpeas The chickpea or chick pea (''Cicer arietinum'') is an annual legume of the family Fabaceae, subfamily Faboideae. Its different types are variously known as gram" or Bengal gram, garbanzo or garbanzo bean, or Egyptian pea. Chickpea seeds are high ...
.
Inga edulis ''Inga edulis'', known as ice-cream bean, ice-cream-bean, joaquiniquil, cuaniquil, guama or guaba, is a fruit native to South America. It is in the Mimosoideae, mimosoid tribe of the legume family Fabaceae. It is widely grown, especially by Indi ...
In
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the Car ...
the
aril An aril (pronounced ), also called an arillus, is a specialized outgrowth from a seed that partly or completely covers the seed. An arillode or false aril is sometimes distinguished: whereas an aril grows from the attachment point of the see ...
s are also used to prepare an
alcoholic beverage An alcoholic beverage (also called an alcoholic drink, adult beverage, or a drink) is a drink that contains ethanol, a type of alcohol that acts as a drug and is produced by fermentation of grains, fruits, or other sources of sugar. The c ...
called cachiri for a festival of the same name. The native women chew the arils and spit the mixture into a vat, where it is left to ferment. The fruit ripens quickly and can only be kept for three to four days, limiting potential export opportunities. However, refrigeration can extend
shelf life Shelf life is the length of time that a commodity may be stored without becoming unfit for use, consumption, or sale. In other words, it might refer to whether a commodity should no longer be on a pantry shelf (unfit for use), or no longer on a ...
to about three weeks. Use in agroforestry systems Due to its rapid growth and ability to improve soil properties, ''Inga edulis'' has found widespread use in tropical
agroforestry Agroforestry is a land use management system in which trees or shrubs are grown around or among crops or pastureland. Trees produce a wide range of useful and marketable products from fruits/nuts, medicines, wood products, etc. This intentional ...
. Most notably,
Inga alley cropping Inga alley cropping is the planting agricultural crops between rows of Inga trees. It has been promoted by Mike Hands. Using the Inga tree for alley cropping has been proposed as an alternative to the much more ecologically destructive slash an ...
is used as an alternative to slash and burn cultivation. It is also a popular shade tree for
coffee Coffee is a drink prepared from roasted coffee beans. Darkly colored, bitter, and slightly acidic, coffee has a stimulant, stimulating effect on humans, primarily due to its caffeine content. It is the most popular hot drink in the world. S ...
,
cocoa Cocoa may refer to: Chocolate * Chocolate * ''Theobroma cacao'', the cocoa tree * Cocoa bean, seed of ''Theobroma cacao'' * Chocolate liquor, or cocoa liquor, pure, liquid chocolate extracted from the cocoa bean, including both cocoa butter and ...
and
tea Tea is an aromatic beverage prepared by pouring hot or boiling water over cured or fresh leaves of '' Camellia sinensis'', an evergreen shrub native to East Asia which probably originated in the borderlands of southwestern China and northe ...
plantations. One important reason for this is that in comparison to other
shade tree A shade tree is a large tree whose primary role is to provide shade in the surrounding environment due to its spreading canopy (forest), canopy and crown (botany), crown, where it may give shelter from sunlight in the heat of the summer for peopl ...
species ''Inga edulis'' retains its leaves in the dry season. Cuttings and leaves also used as mulch and animal fodder. The wood is sometimes used as timber and for fuel, its popularity as a source of firewood stemming from its high calorific value and weak smoke production. Despite its rapid growth and significant potential to improve soils in agroforestry systems, ''Inga edulis'' has not been the object of any improved breeding program. However, the plant has been shown to exhibit strong
introgression Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Intr ...
with ''Inga ingoides'' in species contact areas. This could allow for selection of hybrids via interspecific
hybridization Hybridization (or hybridisation) may refer to: *Hybridization (biology), the process of combining different varieties of organisms to create a hybrid *Orbital hybridization, in chemistry, the mixing of atomic orbitals into new hybrid orbitals *Nu ...
to further increase yield and flood tolerance of the crop. Medicinal uses ''Inga edulis'' is widely used in native South American folk medicine. Decoctions of the leaves, bark or root commonly are utilized as treatments for diarrhea, arthritis and rheumatism. Crushed, boiled leaves are ingested to relieve coughing or applied to lip sores, possibly caused by
herpes Herpes simplex is a viral infection caused by the herpes simplex virus. Infections are categorized based on the part of the body infected. Oral herpes involves the face or mouth. It may result in small blisters in groups often called cold s ...
. Several studies have investigated ''Inga edulis'' as a source of
polyphenols Polyphenols () are a large family of naturally occurring organic compounds characterized by multiples of phenol units. They are abundant in plants and structurally diverse. Polyphenols include flavonoids, tannic acid, and ellagitannin, some of ...
for use as
antioxidant Antioxidants are compounds that inhibit oxidation, a chemical reaction that can produce free radicals. This can lead to polymerization and other chain reactions. They are frequently added to industrial products, such as fuels and lubricant ...
s and have shown promising results. However, further studies are necessary to test and develop medical applications.


References


Ice Cream Beans
* (1983)

Version of 07-JAN-1998. Retrieved 2007-DEC-17. * (2005)
Genus ''Canavalia''
Version 10.01, November 2005. Retrieved 2007-DEC-17. * (2007): Germplasm Resources Information Network
''Inga edulis''
Retrieved 2007-DEC-18. {{Taxonbar, from=Q138183 edulis Edible legumes Flora of South America Tropical fruit Trees of Peru