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''Apios americana'', sometimes called the American groundnut, potato bean, hopniss, Indian potato, hodoimo, America-hodoimo, cinnamon vine, or groundnut (not to be confused with other plants in the subfamily Faboideae sometimes known by that name) is a perennial vine that bears edible
bean A bean is the seed of several plants in the family Fabaceae, which are used as vegetables for human or animal food. They can be cooked in many different ways, including boiling, frying, and baking, and are used in many traditional dishes th ...
s and large edible tubers.


Description

The vine of American groundnut can grow to long. It has pinnate
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
long with 5–7 leaflets. The flowers are usually pink, purple, or red-brown, and are produced in dense
raceme A raceme ( or ) or racemoid is an unbranched, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are produced as the s ...
s in length. The fruit is a
legume 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 f ...
(pod) long. In botanical terms, the tubers are rhizomatous stems, not roots.


Genetics

The species is normally 2n=2x=22,
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
, but both diploid and triploid forms exist. Only diploids are capable of producing seeds; triploids will produce flowers but not seeds. Thus, triploids are entirely dependent on tuber division for propagation whereas diploids can be propagated through both seeds and tubers. Other than seed production, there are no easily identifiable differences between diploids and triploids. Triploids are generally found in the northern part of American groundnut's range whereas diploids predominate in the southern part of the range. Triploids have been identified in Canada (in New Brunswick, Quebec, and Ontario) and the United States (in Connecticut, Vermont, Massachusetts,
New York New York most commonly refers to: * New York City, the most populous city in the United States, located in the state of New York * New York (state), a state in the northeastern United States New York may also refer to: Film and television * '' ...
, Pennsylvania, Ohio, New Hampshire, Rhode Island, Wisconsin, and Iowa). A few diploids have been found in the northeastern part of the range, such as along the Black River in Central Ontario. All samples tested in the Southeastern United States have been found to be diploid.


Distribution and habitat

The plant's natural range is from southern Canada (including Ontario, Quebec, and New Brunswick) down through Florida and west as far as the border of Colorado.


Ecology

The species is a larval host for the ''
Epargyreus clarus ''Epargyreus clarus'', the silver-spotted skipper, is a butterfly of the family Hesperiidae. It is claimed to be the most recognized skipper in North America. occurs in fields, gardens, and at forest edges and ranges from southern Canada thro ...
''.


Nitrogen fixation

American groundnut fixes its own nitrogen, which could be a great advantage in comparison to other roots crops, such as potatoes, true yams, and sweet potatoes. These do not fix their own nitrogen and require large applications of nitrogen fertilizer. American groundnut can be nodulated by bacterial strains that are normally found in symbiosis with soybeans or cowpeas. Research has been done on the potential of the soybean strain '' Bradyrhizobium japonicum'' to nodulate American groundnut. It was found that plants nodulated with ''B. japonicum'' yielded ~30% better than unnodulated plants if no nitrogen fertilizer was used. It was also determined that nodulated plants partitioned more carbon into non-edible shoots when they were given nitrogen fertilizer, whereas unnodulated plants responded to nitrogen fertilizer with greater tuber yields than nodulated plants. This data suggests that nitrogen fertilization may be required to maximize tuber size and yields in ''A. americana''.


Cultivation


Domestication

American groundnut is generally considered to be an undomesticated crop. In her 1939 description of the Native American use of American groundnut, Gretchen Beardsley states that several historical sources describe the "cultivation" of American groundnut by indigenous peoples. She dismisses the ambiguous term "cultivation" as perhaps referring to transplantation of tubers near a settlement. She quotes the historical author Waugh on this subject of cultivation: "sometimes planted in suitable locations, though they are not, strictly speaking, cultivated." Subsequent authors on the American groundnut have followed Beardsley's interpretation of "cultivation" when referring to the early use by Indigenous peoples of American groundnut. However, recent evidence suggests that North American indigenous peoples likely intervened significantly as cultivators of the native plants of the region, in a manner similar to contemporary Western permaculture practices. So, from a permacultural perspective, Native Americans may well have "cultivated" the groundnut. In 1985, Dr. William J. Blackmon, Dr. Berthal D. Reynolds, and their colleagues at Louisiana State University in
Baton Rouge Baton Rouge ( ; ) is a city in and the capital of the U.S. state of Louisiana Louisiana , group=pronunciation (French: ''La Louisiane'') is a state in the Deep South and South Central regions of the United States. It is the 20th-sma ...
, Louisiana, began a program of deliberate domestication of American groundnut. Their primary goal was to develop an American groundnut that can produce a significant yield in a single season. Early trials identified LA85-034 as a promising cultivar, with "elongate tubers of uniform, medium size with light brown skin and little extra rhizomatous material". By 1988, they had collected wild seeds and tubers from 210 plants found in 19 states, although the bulk of their selections came from the state of Louisiana. From these wild materials, and a small number of single crosses, they rigorously selected for plants that met their primary breeding goals of larger tuber size, denser tuber set, single season production, and productivity in untrellised cultivation. The American groundnut domestication program at Louisiana State University continued in various forms until the mid-1990s. Cultivars from this program can still occasionally be found available from small seed companies. From 1985 to 1994, an ''Apios'' breeding program took place that resulted in the collection of over 200 wild accessions. These accessions underwent
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 ...
and selection, and over 2,200 lines were assessed. Of these lines, only 53 genotypes were kept for further analysis. Three different locations and three different growing conditions—field, pots, and grow-bags—were used. There was significant variation found among almost all of the 20 genotypes in the field growing condition. Inter-node length, plant vigor, and stem diameter during plant growth were positively correlated with the plant yield below ground. There were four distinct genotypic clusters found in this collection of ''Apios'' lines. Several genotypes yielded large plants in all locations, maxing up to of subterraneous tuber. This suggests that the plant has a good ability to adapt and grow in a wide variety of locations and conditions. Furthermore, the superior germplasm identified in this project may be suitable as cultivars, and will aid in further development of ''Apios'' lines as a crop. The largest germplasm collection of ''A. americana'' cultivars today is found at Iowa State University under the direction of Dr. Steven Cannon. It is maintained there for scholarly and academic use. Research continues at Iowa State on the domestication of American groundnut. Despite these efforts at domestication, the American groundnut remains largely uncultivated and underused in North America and Europe. There are challenges to breeding and domesticating this plant, as well. There seems to be a partial self-incompatibility with Apios breeding and manual pollinations, resulting in rare seed-sets. Disadvantages in Apios as a crop are its vining habit. The crop has small tuber size for most genotypes. These sizes are typically smaller than ; however, some do average around . The tuber plant is difficult to harvest because of the "beads on a string" arrangement on
stolon In biology, stolons (from Latin '' stolō'', genitive ''stolōnis'' – "branch"), also known as runners, are horizontal connections between organisms. They may be part of the organism, or of its skeleton; typically, animal stolons are external s ...
s, which extend for over a meter.


In Japan and South Korea

The only place in the world today where American groundnuts are commercially farmed in any significant quantities is in
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
. Before the American groundnut was introduced to Japan, the people on the main island of Honshu and the northern island of Hokkaido were already familiar with a native, wild plant called hodoimo (''A. fortunei''), which was occasionally eaten as an emergency food. American groundnut was introduced, accidentally or deliberately, to Japan during the Meiji period (1868–1912). One theory is that it was accidentally brought to Japan as a stowaway weed among apple seedlings imported from North America. Another theory is that American groundnut may have been deliberately brought to Japan in the middle of the Meiji period as an ornamental flower. It has become a culinary specialty of the Aomori Prefecture, where American groundnut agriculture is centered. It has been eaten there for more than one hundred years. Although American groundnut agriculture is primarily identified with agriculture in the Aomori prefecture, it is grown in the nearby prefectures of Akita and Miyagi as well. In addition, it is known to be grown in the southern part of Honshu in the Tottori prefecture. Radioactive testing records following the Fukushima nuclear disaster record cesium testing of American groundnut agricultural products in the central prefecture of Tochigi. An important part of the spread and popularization of American groundnut consumption in Japan has been the efforts of Dr. Kiyochika Hoshikawa to promote the cultivation of this crop in Japan, and the flurry of scientific articles on the health benefits of eating American groundnut tubers. Japanese websites that sell American groundnut continue to emphasize its health benefits in their marketing efforts. There are reports of American groundnut cultivation in South Korea as well, where it is grown for its nutritional benefits.


Toxicity

Studies in rats suggest that raw tubers should not be consumed. They contain harmful protease inhibitors that are denatured by cooking.


Uses

The tubers and seeds can both be cooked and eaten.


By Indigenous peoples of the Americas

The tubers have traditionally been a
staple food A staple food, food staple, or simply a staple, is a food that is eaten often and in such quantities that it constitutes a dominant portion of a standard diet for a given person or group of people, supplying a large fraction of energy needs and ...
among most Indigenous peoples of the Americas within the natural range of the plant. In 1749, the travelling Swedish botanist Peter Kalm writes, "Hopniss or Hapniss was the Indian name of a wild plant, which they ate at that time... The roots resemble potatoes, and were boiled by the Indians who ate them instead of bread." Strachey in 1612 recorded observations of the Indigenous peoples in Virginia: "In June, July, and August they feed upon roots of tockohow, berries, groundnutts, fish, and greene wheate..." In Eastern Canada, the Jesuit missionary, Le Jeune, observed that the Indigenous peoples there would, "eat, besides, roots, such as the bulbs of the red lily; ... another that our French people call 'Rosary' because it is distinguished by tubers in the form of beads." The early author Rafinesque observed that the
Cree The Cree ( cr, néhinaw, script=Latn, , etc.; french: link=no, Cri) are a Indigenous peoples of the Americas, North American Indigenous people. They live primarily in Canada, where they form one of the country's largest First Nations in Canada ...
were cultivating the plant for both its tubers and seeds. The author Brinton wrote in 1885 in regards to the
Lenape The Lenape (, , or Lenape , del, Lënapeyok) also called the Leni Lenape, Lenni Lenape and Delaware people, are an indigenous peoples of the Northeastern Woodlands, who live in the United States and Canada. Their historical territory includ ...
people, "Of wild fruits and plants they consumed the esculent and nutritious tubers on the roots of the Wild Bean, Apios tuberosa... which the Indians called hobbenis..." In 1910, Parker writes that the Iroquois were consuming significant quantities of groundnuts up until about thirty years before his writing. The Paris Documents of 1666 record that the sixth tribe of the second division of the Iroquois were identified as, "that of the Potatoe, which they call Schoneschironon" and an illustration of tubers is found in the Paris Documents with the explanation, "This is the manner they paint the tribe of the Potatoe." The author Gilmore records the use of groundnuts by the Caddoan and Siouan tribes of the Missouri river region, and the authors Prescott and Palmer record its use among the
Sioux The Sioux or Oceti Sakowin (; Dakota language, Dakota: Help:IPA, /otʃʰeːtʰi ʃakoːwĩ/) are groups of Native Americans in the United States, Native American tribes and First Nations in Canada, First Nations peoples in North America. The ...
. The Indigenous peoples would prepare the tubers in many different ways, such as frying them in animal fat or drying them into flour. Many tribes peel them and dry them in the sun, such as the
Menomini The Menominee (; mez, omǣqnomenēwak meaning ''"Menominee People"'', also spelled Menomini, derived from the Ojibwe language word for "Wild Rice People"; known as ''Mamaceqtaw'', "the people", in the Menominee language) are a federally rec ...
who have traditionally built scaffolds of cedar bark covered with mats to dry their tubers for winter use. The Menomini are recorded as having dried the tubers in
maple syrup Maple syrup is a syrup made from the sap of maple trees. In cold climates, these trees store starch in their trunks and roots before winter; the starch is then converted to sugar that rises in the sap in late winter and early spring. Maple tree ...
or making a preserve of Groundnut tubers by boiling them in maple syrup. The
Potawatomi The Potawatomi , also spelled Pottawatomi and Pottawatomie (among many variations), are a Native American people of the western Great Lakes region, upper Mississippi River and Great Plains. They traditionally speak the Potawatomi language, a m ...
have traditionally boiled their tubers. The traditional Meskwaki and Chippewa preparation involves peeling, parboiling, slicing, and drying the tubers. The Chippewa have historically used them as a sort of seasoning in all their foods.


By Europeans

The Europeans learned to use the American groundnut from the Indigenous peoples of the Americas. As a result, the American groundnut became interwoven with the history of the American colonies and Europe. The early traveler John Brereton was sustained by the "good meat" and "medicinable" qualities of American groundnut during his travels in New England in 1602. In 1613, the followers of Biencourt at Port-Royal ate the tubers to help them survive in the New World. The American groundnut was an important factor in the survival of the Pilgrims during the first few winters of their settlement. In 1623 the Pilgrims, "having but a small quantity of corn left," were "enforced to live on groundnuts... and such other things that the country afforded... and were easily gotten..." The Pilgrims were taught to find and prepare American groundnut by the
Wampanoag people The Wampanoag , also rendered Wôpanâak, are an Indigenous people of the Northeastern Woodlands based in southeastern Massachusetts and historically parts of eastern Rhode Island,Salwen, "Indians of Southern New England and Long Island," p. 17 ...
. The groundnut was likely eaten at the harvest festival of November 1621 that is regarded as the first Thanksgiving, although only venison was specifically named as a food item at this meal by a Pilgrim eyewitness account. Philosopher
Henry David Thoreau Henry David Thoreau (July 12, 1817May 6, 1862) was an American naturalist, essayist, poet, and philosopher. A leading Transcendentalism, transcendentalist, he is best known for his book ''Walden'', a reflection upon simple living in natural su ...
commented on the nutty flavor and dry texture in October 1852. It is believed that American groundnut may have been shipped to Europe as early as 1597. It was listed in 1885 as a European garden crop. In 1845 it was evaluated as a possible alternative potato crop in Ireland during the Great Famine. These early introductions to Europe appear to have resulted in little or no assimilation of the new food into the European diet. A primary reason for this lack of assimilation was that the two-year cycle for an acceptable tuber yield did not match the cropping systems that were familiar to Europeans.


Nutrition

The tubers are highly palatable with culinary characteristics of a potato, although the flavor can be somewhat nuttier than a potato and the texture can be finer. These tubers contain roughly three times the protein content of a potato (16.5% by dry weight), and the amino acid balance is good with the exception of
cysteine Cysteine (symbol Cys or C; ) is a semiessential proteinogenic amino acid with the formula . The thiol side chain in cysteine often participates in enzymatic reactions as a nucleophile. When present as a deprotonated catalytic residue, sometime ...
and
methionine Methionine (symbol Met or M) () is an essential amino acid in humans. As the precursor of other amino acids such as cysteine and taurine, versatile compounds such as SAM-e, and the important antioxidant glutathione, methionine plays a critical ro ...
. ''A. americana'' tubers were found to have a protein concentration of , similar to that of other species in the genus, '' A. carnea'' and '' A. fortunei''. However, ''A. americana'' had larger levels of genistein than the other two species. The fatty acid content of tubers is approximately 4.2% to 4.6%, with linoleic fatty acids predominating. The fresh weight of a tuber is 36% of carbohydrate (primarily starch). The tubers are also an excellent source of calcium and iron. Calcium content is tenfold greater than a potato and iron is twofold greater than a potato, although vitamin C was considerably less than a potato. The tuber and the flower also contain
monosaccharide Monosaccharides (from Greek ''monos'': single, '' sacchar'': sugar), also called simple sugars, are the simplest forms of sugar and the most basic units (monomers) from which all carbohydrates are built. They are usually colorless, water-solub ...
s and oligosaccharides. The tuber has more of these types of carbohydrates than the soybean, potato, and
sweet potato The sweet potato or sweetpotato (''Ipomoea batatas'') is a dicotyledonous plant that belongs to the Convolvulus, bindweed or morning glory family (biology), family, Convolvulaceae. Its large, starchy, sweet-tasting tuberous roots are used as a r ...
. In addition, the tubers appear to have numerous health-promoting factors. Hypertensive rats that were fed powdered tubers as 5% of their total diet had a 10% decrease in blood pressure and also a reduction in cholesterol and triglycerides. It has been shown that the tubers contain genistein and other
isoflavones Isoflavones are substituted derivatives of isoflavone, a type of naturally occurring isoflavonoids, many of which act as phytoestrogens in mammals. Isoflavones are produced almost exclusively by the members of the bean family, Fabaceae (Leguminosae) ...
that have various health benefits, including an anti-carcinogenic function against colon, prostate, and breast cancer. Genistein-7-O-gentiobioside is a novel isoflavone that is found in the American groundnut. Extract from the American groundnut was shown to drive the anti-oxidative pathway in cells although it did not have anti-oxidative activity itself. Human breast carcinoma MCF-7 cells were pretreated with the extract of ''A. americana'' for 24 hours. Subsequent analysis showed an increase in expression of
heme oxygenase-1 Heme oxygenase, or haem oxygenase, (HMOX, commonly abbreviated as HO) is an enzyme that catalyzes the degradation of heme to produce biliverdin, ferrous ion, and carbon monoxide. There are many heme degrading enzymes in nature. In general, only ...
, a protein induced during oxidative stress. The American groundnut, like soybean, is a great source of isoflavone. Furthermore, a study on ''A. americana'' and its flower shows that the flower of the particular plant is not toxic to mice. Consumption of the flower was shown to lower plasma glucose levels in diabetic mice. The flower was shown to have an inhibitory activity on
maltose } Maltose ( or ), also known as maltobiose or malt sugar, is a disaccharide formed from two units of glucose joined with an α(1→4) bond. In the isomer isomaltose, the two glucose molecules are joined with an α(1→6) bond. Maltose is the two- ...
and an anti-hyperglycemic effect in mice, suggesting that not only is it a viable and novel food source for the general population, but also in the prevention of diabetes.


References


External links


Domestication of ''Apios americana''Plants for a future database''Apios americana'' at the USDA Plants databaseOrion magazine articleWired magazine - How We Can Tame Overlooked Wild Plants to Feed the World
{{Taxonbar, from=Q568933 Phaseoleae Edible legumes Plants used in Native American cuisine Root vegetables Perennial vegetables Flora of the Northeastern United States Flora of the Southeastern United States Flora of the Great Lakes region (North America) Flora of the United States Plants described in 1753 Taxa named by Carl Linnaeus Flora without expected TNC conservation status