Sorghum

Sorghum bicolor, commonly called sorghum[2] (/ˈsɔːrɡəm/) and also known as great millet,[3] broomcorn,[4] guinea corn,[5] durra,[6] imphee,[7] jowar,[8] or milo,[9] is a grass species cultivated for its grain, which is used for food for humans, animal feed, and ethanol production. Sorghum originated in Africa, and is now cultivated widely in tropical and subtropical regions.[10]

Sorghum
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Clade: Commelinids
Order: Poales
Family: Poaceae
Subfamily: Panicoideae
Genus: Sorghum
Species:
S. bicolor
Binomial name
Sorghum bicolor
(L.) Moench
Synonyms[1]
List
    • Agrostis nigricans (Ruiz & Pav.) Poir.
    • Andropogon besseri Kunth
    • Andropogon bicolor (L.) Roxb.
    • Andropogon caffrorum (Thunb.) Kunth
    • Andropogon compactus Brot.
    • Andropogon dulcis Burm.f.
    • Andropogon niger (Ard.) Kunth
    • Andropogon saccharatrus Kunth
    • Andropogon saccharatus (L.) Raspail
    • Andropogon sorghum (L.) Brot.
    • Andropogon subglabrescens Steud.
    • Andropogon truchmenorum Walp.
    • Andropogon usorum Steud.
    • Andropogon vulgare (Pers.) Balansa
    • Andropogon vulgaris Raspail
    • Holcus arduinii J.F.Gmel.
    • Holcus bicolor L.
    • Holcus cafer Ard.
    • Holcus caffrorum (Retz.) Thunb.
    • Holcus cernuus Ard.
    • Holcus cernuus Muhl. nom. illeg.
    • Holcus cernuus Willd. nom. illeg.
    • Holcus compactus Lam.
    • Holcus dochna Forssk.
    • Holcus dora Mieg
    • Holcus duna J.F.Gmel.
    • Holcus durra Forssk.
    • Holcus niger Ard.
    • Holcus nigerrimus Ard.
    • Holcus rubens Gaertn.
    • Holcus saccharatus var. technicus (Körn.) Farw.
    • Holcus sorghum L.
    • Holcus sorghum Brot. nom. illeg.
    • Milium bicolor (L.) Cav.
    • Milium compactum (Lam.) Cav.
    • Milium maximum Cav.
    • Milium nigricans Ruiz & Pav.
    • Milium sorghum (L.) Cav.
    • Panicum caffrorum Retz.
    • Panicum frumentaceum Salisb. nom. illeg.
    • Rhaphis sorghum (L.) Roberty
    • Sorghum abyssinicum (Hack.) Chiov. nom. illeg.
    • Sorghum ankolib (Hack.) Stapf
    • Sorghum anomalum Desv.
    • Sorghum arduinii (Gmel.) J.Jacq.
    • Sorghum basiplicatum Chiov.
    • Sorghum basutorum Snowden
    • Sorghum caffrorum (Retz.) P.Beauv.
    • Sorghum campanum Ten. & Guss.
    • Sorghum caudatum (Hack.) Stapf
    • Sorghum centroplicatum Chiov.
    • Sorghum cernuum (Ard.) Host
    • Sorghum compactum Lag.
    • Sorghum conspicuum Snowden
    • Sorghum coriaceum Snowden
    • Sorghum dochna (Forssk.) Snowden
    • Sorghum dora (Mieg) Cuoco
    • Sorghum dulcicaule Snowden
    • Sorghum dura Griseb.
    • Sorghum durra (Forssk.) Batt. & Trab.
    • Sorghum elegans (Körn.) Snowden
    • Sorghum eplicatum Chiov.
    • Sorghum exsertum Snowden
    • Sorghum gambicum Snowden
    • Sorghum giganteum Edgew.
    • Sorghum glabrescens (Steud.) Schweinf. & Asch.
    • Sorghum glycychylum Pass.
    • Sorghum guineense Stapf
    • Sorghum japonicum (Hack.) Roshev.
    • Sorghum margaritiferum Stapf
    • Sorghum medioplicatum Chiov.
    • Sorghum melaleucum Stapf
    • Sorghum melanocarpum Huber
    • Sorghum mellitum Snowden
    • Sorghum membranaceum Chiov.
    • Sorghum miliiforme (Hack.) Snowden
    • Sorghum nankinense Huber
    • Sorghum nervosum Besser ex Schult. & Schult.f.
    • Sorghum nervosum Chiov. nom. illeg.
    • Sorghum nigricans (Ruiz & Pav.) Snowden
    • Sorghum nigrum (Ard.) Roem. & Schult.
    • Sorghum notabile Snowden
    • Sorghum pallidum Chiov. nom. illeg.
    • Sorghum papyrascens Stapf
    • Sorghum rigidum Snowden
    • Sorghum rollii Chiov.
    • Sorghum roxburghii var. hians (Hook.f.) Stapf
    • Sorghum saccharatum Host nom. illeg.
    • Sorghum saccharatum (L.) Pers. nom. illeg.
    • Sorghum sativum (Hack.) Batt. & Trab.
    • Sorghum schimperi (Hack.) Chiov. nom. illeg.
    • Sorghum simulans Snowden
    • Sorghum splendidum (Hack.) Snowden
    • Sorghum subglabrescens (Steud.) Schweinf. & Asch.
    • Sorghum tataricum Huber
    • Sorghum technicum (Körn.) Batt. & Trab.
    • Sorghum technicum (Körn.) Roshev.
    • Sorghum truchmenorum K.Koch
    • Sorghum usorum Nees
    • Sorghum vulgare Pers. nom. illeg.

Sorghum is the world's fifth-most important cereal crop after rice, wheat, maize, and barley, with 61,000,000 metric tons (60,000,000 long tons; 67,000,000 short tons) of annual global production in 2021.[11] Sorghum is typically an annual, but some cultivars are perennial. It grows in clumps that may reach over 4 metres (13 ft) high. The grain is small, ranging from 2 to 4 millimetres (0.079 to 0.157 in) in diameter. Sweet sorghums are cultivars primarily grown for forage, syrup production, and ethanol; they are taller than those grown for grain.[12][13]

Sorghum bicolor is the cultivated species of sorghum; its wild relatives make up the botanical genus Sorghum.

Description

Sorghum is a large stout grass that grows up to 2.4 metres (7.9 ft) tall. It has large bushy flowerheads or panicles that provide an edible starchy grain with up to 3,000 seeds in each flowerhead. It grows in warm climates worldwide.[14][15] One species, S. bicolor, is grown worldwide in large quantities for food and forage.[16] native to Africa with many cultivated forms.[17] Seventeen of the 25 species are native to Australia,[18][19][20] with the range of some extending to Africa, Asia, Mesoamerica, and certain islands in the Indian and Pacific Oceans.[21][22]

Evolution

Phylogeny

Sorghum is closely related to maize within the PACMAD clade of grasses, and more distantly to the cereals of the BOP clade such as wheat and barley.[23]

(Part of Poaceae)
BOP clade

Bambusoideae (bamboos)

Pooideae
other grasses

 (fescue, ryegrass)

  Triticeae  

Hordeum (barley)

Triticum (wheat)

Secale (rye)

Oryza (rice)

PACMAD clade

Pennisetum (fountaingrasses, pearl millet)

Sorghum (sorghum)

Zea (maize)

Taxonomy

Sorghum is in the grass family, Poaceae, in the subfamily Panicoideae, in the tribe Andropogoneae the same as maize (Zea mays), big bluestem (Andropogon gerardi), and sugarcane (Saccharum spp.).

Accepted species recorded include:[24]

Sorghum flour on sale in a street market, Ethiopia, 2013
A plate of sorghum grain
  • Sorghum amplum – northwestern Australia
  • Sorghum angustum – Queensland
  • Sorghum arundinaceum – Africa, Indian Subcontinent, Madagascar, islands of the western Indian Ocean
  • Sorghum bicolor – cultivated sorghum, often individually called sorghum, also known as durra, jowari, or milo. Native to Sahel region of Africa; naturalized in many places
  • Sorghum brachypodum – Northern Territory of Australia
  • Sorghum bulbosum – Northern Territory, Western Australia
  • Sorghum burmahicum – Thailand, Myanmar
  • Sorghum controversum – India
  • Sorghum × drummondii – Sahel and West Africa
  • Sorghum ecarinatum – Northern Territory, Western Australia
  • Sorghum exstans – Northern Territory of Australia
  • Sorghum grande – Northern Territory, Queensland
  • Sorghum halepense – Johnson grass – North Africa, islands of eastern Atlantic, southern Asia from Lebanon to Vietnam; naturalized in East Asia, Australia, the Americas
  • Sorghum interjectum – Northern Territory, Western Australia
  • Sorghum intrans – Northern Territory, Western Australia
  • Sorghum laxiflorum – Philippines, Lesser Sunda Islands, Sulawesi, New Guinea, northern Australia
  • Sorghum leiocladum – Queensland, New South Wales, Victoria
  • Sorghum macrospermum – Northern Territory of Australia
  • Sorghum matarankense – Northern Territory, Western Australia
  • Sorghum nitidum – East Asia, Indian Subcontinent, Southeast Asia, New Guinea, Micronesia
  • Sorghum plumosum – Australia, New Guinea, Indonesia
  • Sorghum propinquum – China, Indian Subcontinent, Southeast Asia, New Guinea, Christmas Island, Micronesia, Cook Islands
  • Sorghum purpureosericeum – Sahel from Mali to Tanzania; Yemen, Oman, India
  • Sorghum stipoideum – Northern Territory, Western Australia
  • Sorghum timorense – Lesser Sunda Islands, Maluku, New Guinea, northern Australia
  • Sorghum trichocladum – Mexico, Guatemala, Honduras
  • Sorghum versicolor – eastern + southern Africa from Ethiopia to Namibia; Oman
  • Sorghum virgatum – dry regions from Senegal to the Levant.

History

Domestication

Piece of sorghum bread contained in basket, Predynastic Egypt, c. 3100 BC (5,100 years ago). Egyptian Museum, Turin[25]

S. bicolor was domesticated from its wild ancestor more than 5,000 years ago in Eastern Sudan in the area of the Rivers Atbara and Gash.[26][27] It has been found at an archaeological site near Kassala in eastern Sudan, dating from 3500 to 3000 BC, and is associated with the neolithic Butana Group culture.[28] Sorghum bread from graves in Predynastic Egypt, some 5,100 years ago, is displayed in the Egyptian Museum, Turin, Italy.[25]

The first race to be domesticated was bicolor; it had tight husks that had to be removed forcibly. Around 4,000 years ago, this spread to the Indian subcontinent; around 3,000 years ago it reached West Africa.[26] Four other races evolved through cultivation to have larger grains and to become free-threshing, making harvests easier and more productive. These were caudatum in the Sahel; durra, most likely in India; guinea in West Africa (later reaching India), and from that race mageritiferum that gave rise to the varieties of Southern Africa.[26]

Domestication and races of sorghum

Spread

In the Middle Ages, the Arab Agricultural Revolution spread sorghum and other crops from Africa and Asia across the Arab world as far as Al-Andalus in Spain.[29] Sorghum remained the staple food of the medieval kingdom of Alodia and most Sub-Saharan cultures prior to European colonialism.[30]

Sweet sorghum was important to the sugar trade in the 19th century.[31] The price of sugar was rising because of decreased production in the British West Indies and more demand for confectionery and fruit preserves, and the United States was actively searching for a sugar plant that could be produced in northern states. The "Chinese sugar-cane", sweet sorghum, was viewed as a plant that would be productive in the West Indies.[32]

Nutrition

Sorghum grain
Nutritional value per 100 g (3.5 oz)
Energy329 kJ (79 kcal)
72.1 g
Sugars2.53 g
Dietary fiber6.7 g
3.46 g
Saturated0.61 g
Monounsaturated1.13 g
Polyunsaturated1.56 g
10.6 g
VitaminsQuantity
%DV
Vitamin A equiv.
0%
0 μg
Thiamine (B1)
28%
0.332 mg
Riboflavin (B2)
7%
0.096 mg
Niacin (B3)
23%
3.69 mg
Pantothenic acid (B5)
7%
0.367 mg
Vitamin B6
26%
0.443 mg
Folate (B9)
5%
20 μg
Vitamin C
0%
0 mg
Vitamin E
3%
0.5 mg
MineralsQuantity
%DV
Calcium
1%
13 mg
Copper
32%
0.284 mg
Iron
19%
3.36 mg
Magnesium
39%
165 mg
Manganese
70%
1.6 mg
Phosphorus
23%
289 mg
Potassium
12%
363 mg
Selenium
22%
12.2 μg
Sodium
0%
2 mg
Zinc
15%
1.67 mg
Other constituentsQuantity
Water12.4 g

Percentages estimated using US recommendations for adults,[33] except for potassium, which is estimated based on expert recommendation from the National Academies.[34]

The grain is edible and nutritious. It can be eaten raw when young and milky, but has to be boiled or ground into flour when mature.[35]

Sorghum grain is 72% carbohydrates including 7% dietary fiber, 11% protein, 3% fat, and 12% water (table). In a reference amount of 100 grams (3.5 oz), sorghum grain supplies 79 calories and rich contents (20% or more of the Daily Value, DV) of several B vitamins and dietary minerals (table).

In the early stages of plant growth, some sorghum species may contain levels of hydrogen cyanide, hordenine, and nitrates lethal to grazing animals.[36] Plants stressed by drought or heat can also contain toxic levels of cyanide and nitrates at later stages in growth.[37]

Use

S. bicolor is widely used for food and animal fodder. It is also used to make alcoholic beverages, and biofuels such as ethanol.[15] It can be made into couscous, porridge, or flatbreads such as Indian Jōḷada roṭṭi or tortillas; and it can be burst in hot oil to make a popcorn, smaller than that of maize. Since it does not contain gluten, it can be used in gluten-free diets.[38]

In Nigeria, the pulverized red leaf-sheaths of sorghum have been used to dye leather, while in Algeria, sorghum has been used to dye wool.[39]

In South Africa, characteristically sour Kaffir beer is made from sorghum or millet. The process involves souring the mashed grain with lactic acid bacteria, followed by fermenting by the wild yeasts that were on the grain.[40]

In India, the panicle stalks are used as bristles for brooms.[41]

In countries including the US, the stalks of sweet sorghum varieties are crushed in a cane juicer to extract the sweet molasses-like juice. The juice is sold as syrup,[42][43][44] and used as a feedstock to make biofuel.[45]

Cultivation

Agronomy

Most varieties of sorghum are drought- and heat-tolerant, nitrogen-efficient,[46] and are grown particularly in arid and semi-arid regions where the grain is one of the staples for poor and rural people. These varieties provide forage in many tropical regions. S. bicolor is a food crop in Africa, Central America, and South Asia, and is the fifth most common cereal crop grown in the world.[47][48] It is most often grown without application of fertilizers or other inputs by small-holder farmers in developing countries.[49]

Sorghum grows in a wide range of temperatures. It can tolerate high altitude and toxic soils, and can recover growth after some drought.[12] Optimum growth temperature range is 12–34 °C (54–93 °F), and the growing season lasts for ~ 115-140 days. [50] It can grow on a wide range of soils, such as heavy clay to sandy soils with the pH tolerance ranging from 5.0 to 8.5.[51] It requires an arable field that has been left fallow for at least two years or where crop rotation with legumes has taken place in the previous year.[52] Diversified 2- or 4-year crop rotation can improve sorghum yield, additionally making it more resilient to inconsistent growth conditions.[53] In terms of nutrient requirements, sorghum is comparable to other cereal grain crops with nitrogen, phosphorus, and potassium required for growth.[54]

Pests and diseases

Insect damage is a major threat to sorghum plants. Over 150 species damage crop plants at different stages of development, resulting in significant biomass loss.[55] Stored sorghum grain is attacked by other insect pests such as the lesser grain borer beetle.[56] Sorghum is a host of the parasitic plant Striga hermonthica, purple witchweed; it can seriously reduce production.[57] Sorghum is subject to a variety of plant pathogens. The fungus Colletotrichum sublineolum causes anthracnose.[58] The toxic ergot fungus parasitises the grain, risking harm to humans and livestock.[59] Sorghum produces chitinases as defensive compounds against fungal diseases. Transgenesis of additional chitinases increases the crop's disease resistance.[60]

Genetics and genomics


The genome of S. bicolor was sequenced between 2005 and 2007.[61][62] It is generally considered diploid and contains 20 chromosomes,[63] however, there is evidence to suggest a tetraploid origin for S. bicolor.[64] The genome size is approximately 800 Mbp.[65]

Paterson et al., 2009 provides a genome assembly of 739 megabase. The most commonly used genome database is SorGSD maintained by Luo et al., 2016. A gene expression atlas is available from Shakoor et al., 2014 with 27,577 genes. For molecular breeding (or other purposes) an SNP array has been created by Bekele et al., 2013, a 3K SNP Infinium from Illumina, Inc.[66]

Agrobacterium transformation can be used on sorghum, as shown in a 2018 report of such a transformation system.[67] A 2013 study developed and validated an SNP array for molecular breeding.[66][68]

Production

Sorghum production – 2021
Country (Millions of tonnes)
 United States11.4
 India4.8
 Ethiopia4.4
 Mexico4.4
 Argentina3.3
 China3.0
World61.4
Source: FAOSTAT of the United Nations[69]

In 2021, world production of sorghum was 61 million tonnes, led by the United States with 19% of the total (table). India, Ethiopia, and Mexico were the largest secondary producers.[69]

International trade

In 2013, China began purchasing American sorghum as a complementary livestock feed to its domestically grown maize. It imported around $1 billion worth per year until April 2018, when it imposed retaliatory tariffs as part of a trade war.[70] By 2020, the tariffs have been waived, and trade volumes increased again[71] before declining again as China began buying sorghum from other countries.[72] As of 2020, China is the world's largest sorghum importer, importing more than all other countries combined.[71]

In human culture

In Australia, sorghum is personified as a spirit among the Dagoman people of Northern Territory, as well as being used for food; the local species are S. intrans and S. plumosum.[73]

In Korea, the origin tale "Brother and sister who became the Sun and Moon" is also called "The reason sorghum is red".[74] In the tale, a tiger who is chasing a brother and sister follows them up a rotten rope as they climb into the sky, and become the sun and moon. The rope breaks, and the tiger falls to its death, impaling itself on a sorghum stalk, which becomes red with its blood.[75]

In Northeastern Italy in the early modern period, sticks of sorghum were used by Benandanti visionaries of the Friuli district to fight off witches who were thought to threaten crops and people.[76]

See also

References

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