Biology:Sorghum

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Sorghum bicolor, commonly called sorghum[1] (/ˈsɔːrɡəm/) and also known as broomcorn,[2] great millet,[3] Indian millet,[4] Guinea corn,[5] jowar,[6] or milo,[7] is a species in the grass genus Sorghum. It 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 2 to 4 millimetres (0.08 to 0.2 in) in diameter.

The species originated and was domesticated in Sudan. Native to Africa and the Indian subcontinent, it is cultivated in tropical and subtropical regions for its grain. It is the world's fifth-most important cereal crop. The grain is used as food by humans, while the plant is used for animal feed and ethanol production.

Description

Sorghum is a large stout grass that grows up to 2.4 metres (8 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 for food and forage.[8][9][10] Sorghum is native to Africa and to the Indian Subcontinent, with many cultivated forms.[11][12][13] Most production uses annual cultivars, but some wild species of Sorghum are perennial; the Land Institute is attempting to develop a perennial cultivar for "repeated, sufficient grain harvests without resowing".[14][15] The name sorghum derives from Italian sorgo, which in turn most likely comes from 12th century Medieval Latin surgum or suricum. This in turn may be from Latin syricum, meaning "[grass] of Syria".[16]

Types include milo,[17] durra,[18] imphee,[19] hegari,[20] kaffir, feterita, shallu, and kaoliang.[21]

Evolution

Phylogeny

Sorghum is closely related to maize and the millets within the PACMAD clade of grasses, and more distantly to the cereals of the BOP clade such as wheat and barley.[22]

(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)

 
 
 

Millets

 
 
 

Sorghum (sorghum)

 
 

Zea (maize)

 
 
 
 
 

History

Domestication

File:Piece of sorghum bread contained in basket S - Museo Egizio, Turin S 285 p09.jpg
Piece of sorghum bread contained in basket, Predynastic Egypt, c. 3100 BC (5,100 years ago). Egyptian Museum, Turin[23]

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.[24][25][26] It has been found at an archaeological site near Kassala in eastern Sudan, dating from 3500 to 3000 BCE, and is associated with the Neolithic Butana Group culture.[27] Sorghum bread from graves in Predynastic Egypt, some 5,100 years ago, is displayed in the Egyptian Museum, Turin, Italy.[23]

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.[24] 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 magentiferum that gave rise to the varieties of Southern Africa.[24] Wetter conditions in parts of the Horn of Africa between about 2,500 and 1,000 years ago supported the expansion of sorghum cultivation and contributed to the development of complex agricultural societies such as Aksum.[28]

File:Domestication and races of Sorghum.svg
Domestication and the five major races of sorghum[24]

Spread

File:"Another view of the making of sorghum molasses on the Fred Hatmaker farm in the Norris Dam reservoir area." - NARA - 532657.tif

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]

Tall varieties of sorghum with a high sugar content are called sweet sorghum; these are useful for producing a sugar-rich syrup and as forage.[31][32] Sweet sorghum was important to the sugar trade in the 19th century.[33] 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.[34]

Cultivation

Agronomy

Most varieties of sorghum are drought- and heat-tolerant, nitrogen-efficient,[35] 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.[36][37] It is usually grown without fertilizers or other inputs by small-holder farmers in developing countries.[38] They benefit from sorghum's ability to compete effectively with weeds, especially when planted in narrow rows. Sorghum actively suppresses weeds by producing sorgoleone, an alkylresorcinol.[39]

Sorghum grows in a wide range of temperatures. It can tolerate high altitude and toxic soils, and can recover growth after some drought.[31] Optimum growth temperature range is 12–34 °C (54–93 °F), and the growing season lasts for around 115–140 days.[40] It can grow in a wide range of soils, such as heavy clay to sandy soils with the pH tolerance ranging from 5.0 to 8.5.[41] 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.[42] Diversified 2- or 4-year crop rotation can improve sorghum yield, making it more resilient to inconsistent growth conditions.[43] Nutrients required by sorghum are comparable to other cereal grain crops with nitrogen, phosphorus, and potassium needed for growth.[44]

The International Crops Research Institute for the Semi-Arid Tropics has improved sorghum using traditional genetic improvement and integrated genetic and natural resources management practices.[45] Some 194 improved cultivars are planted worldwide.[46] In India, increases in sorghum productivity resulting from improved cultivars have freed up 7 million hectares (17 million acres) of land, enabling farmers to diversify into high-income cash crops and boost their livelihoods.[47] Sorghum is used primarily as poultry feed, and also as cattle feed and in brewing.[48]

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.[49] Stored sorghum grain is attacked by insect pests such as the lesser grain borer beetle.[50] Sorghum is a host of the parasitic plant Striga hermonthica, purple witchweed that can reduce production.[51] Sorghum is subject to a variety of plant pathogens. The fungus Colletotrichum sublineolum causes anthracnose.[52] The toxic ergot fungus attacks the grain, risking harm to humans and livestock.[53] Sorghum produces chitinases as defensive compounds against fungal diseases. Transgenesis of additional chitinases increases the crop's disease resistance.[54]

Genetics and genomics

The genome of S. bicolor was sequenced between 2005 and 2007.[55][56] It is generally considered diploid and contains 20 chromosomes,[57] however, there is evidence to suggest a tetraploid origin for S. bicolor.[58] The genome size is approximately 800 Mbp.[59]

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.[60]

Agrobacterium transformation can be used on sorghum, as shown in a 2018 report of such a transformation system.[61] A 2013 study developed and validated an SNP array for molecular breeding.[60][62]

Production

Sorghum production
2023, million tonnes
Country Volume
 United States 8.1
 Mexico 4.8
 Ethiopia 4.0
 India 3.8
 China 3.0
 Argentina 1.6
World 57.3
Source: FAOSTAT, United Nations[63]

In 2023, world production of sorghum was 57 million tonnes, led by the United States with 14% of the total (table). Mexico, Ethiopia, and India were secondary producers.[63]

It is the world's fifth-most important cereal crop after rice, wheat, maize, and barley.[64]

Sorghum-growing areas of the US, the world's largest producer

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.[65] By 2020, the tariffs had been waived, and trade volumes increased [66] before declining again as China began buying sorghum from other countries.[67] As of 2020, China is the world's largest sorghum importer, importing more than all other countries combined.[66] Mexico also accounts for 7% of global sorghum production.[68]

Nutrition

Sorghum grain
Nutritional value per 100 g (3.5 oz)
Energy1,380 kJ (330 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)
29%
0.332 mg
Riboflavin (B2)
8%
0.096 mg
Niacin (B3)
25%
3.69 mg
Pantothenic acid (B5)
7%
0.367 mg
Vitamin B6
34%
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
14%
0.284 mg
Iron
26%
3.36 mg
Magnesium
46%
165 mg
Manganese
76%
1.6 mg
Phosphorus
41%
289 mg
Potassium
8%
363 mg
Selenium
17%
12.2 μg
Sodium
0%
2 mg
Zinc
18%
1.67 mg
Other constituentsQuantity
Water12.4 g
Link to USDA Database entry
†Percentages are roughly approximated using US recommendations for adults.
Source: USDA Nutrient Database

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.[69]

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 329 calories and rich contents (20% or more of the Daily Value, DV) of several B vitamins and dietary minerals (table).[70]

In the early stages of plant growth, some sorghum species may contain levels of hydrogen cyanide, hordenine, and nitrates lethal to grazing animals.[71] Plants stressed by drought or heat can also contain toxic levels of cyanide and nitrates at later stages in growth.[72]

Use

Food and drink

Sorghum is widely used for food and animal fodder. It is also used to make alcoholic beverages.[9] 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.[73]

In South Africa, characteristically sour malwa 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.[74] In China and Taiwan, sorghum is one of the main materials of Kaoliang liquor, a type of the colourless distilled alcoholic drink baijiu.[75][76]

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,[77][78][79] and used as a feedstock to make biofuel.[80]

Biofuel

Sorghum can be used to produce fuel ethanol as an alternative to maize. The energy ratio for the production of ethanol is similar to that of sugarcane, and much higher than that of maize.[81] Extracted carbohydrates can readily be fermented into ethanol because of their simple sugar structure. Residuals contain enough energy to be burned to power the ethanol processing facilities used to produce the fuel.[82] As of 2018, production costs (including price of produce,[83] transport and processing costs) are competitive with maize,[84] while sorghum has a lower nitrogen fertilizer requirement than maize.[85] To turn it into fuel ethanol, sorghum juice is concentrated into syrup for long term storage, then fermented in a batch fermentation process.[86]

The stalk of sweet sorghum varieties, called sorgo or sorgho and taller than those grown for grain, can be used as forage or silage and to produce ethanol.[87]

Other uses

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.[88]

In India, the panicle stalks are used as bristles for brooms.[89]

Sorghum seeds and bagasse have the potential to produce lactic acid via fermentation which can be used to make polylactic acid, a biodegradable thermoplastic resin.[90]

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.[91]

In Korea, the origin tale "Brother and sister who became the Sun and Moon" is also called "The reason sorghum is red".[92] 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.[93]

In Northeastern Italy in the early modern period, stalks of fennel were used by Benandanti visionaries, or out-of-body travelers, of the Friuli district to fight off Maledanti, who fought with sticks of sorghum, and were thought to threaten crops and people.[94]

See also

References

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  2. ↑ "Definition of Broomcorn". https://www.merriam-webster.com/dictionary/broomcorn. 
  3. ↑ (xls) BSBI List 2007, Botanical Society of Britain and Ireland, https://bsbi.org/download/3542/, retrieved 14 December 2021 
  4. ↑ Worth, Tammy (28 March 2024). "Sorghum: Are There Health Benefits?". WebMD Health Corp.. https://www.webmd.com/diet/health-benefits-sorghum. 
  5. ↑ "Definition of Guinea corn". https://www.merriam-webster.com/dictionary/Guinea+corn. 
  6. ↑ "jowar". The Free Dictionary. https://www.thefreedictionary.com/jowar. 
  7. ↑ "Milo, Grain Sorghum | Grain Crops". https://graincrops.mgcafe.uky.edu/milo-grain-sorghum. 
  8. ↑ "Sorghum", County-level distribution maps from the North American Plant Atlas (NAPA) (Biota of North America Program (BONAP)), 2014, http://bonap.net/NAPA/TaxonMaps/Genus/County/Sorghum, retrieved 4 September 2016 
  9. ↑ 9.0 9.1 "sorghum: grain". Britannica. https://www.britannica.com/plant/sorghum-grain. 
  10. ↑ Mutegi, Evans; Sagnard, Fabrice; Muraya, Moses; Kanyenji, Ben; Rono, Bernard et al. (2010-02-01). "Ecogeographical distribution of wild, weedy and cultivated Sorghum bicolor (L.) Moench in Kenya: implications for conservation and crop-to-wild gene flow". Genetic Resources and Crop Evolution 57 (2): 243–253. doi:10.1007/s10722-009-9466-7. Bibcode: 2010GRCEv..57..243M. http://oar.icrisat.org/2032/1/GRCE57_243-253__2010.pdf. 
  11. ↑ Hauser, Stefan; Wairegi, Lydia; Asadu, Charles L.A.; Asawalam, Damian O.; Jokthan, Grace; Ugbe, Utiang (2015). "Sorghum- and millet-legume cropping systems". Centre for Agriculture and Bioscience International and Africa Soil Health Consortium. http://africasoilhealth.cabi.org/wpcms/wp-content/uploads/2015/03/392-ASHC-English-Sorghum-BW-A4-lowres.pdf. 
  12. ↑ Dillon, Sally L.; Shapter, Frances M.; Henry, Robert J. et al. (1 September 2007). "Domestication to Crop Improvement: Genetic Resources for Sorghum and Saccharum (Andropogoneae)". Annals of Botany 100 (5): 975–989. doi:10.1093/aob/mcm192. PMID 17766842. 
  13. ↑ "Sorghum bicolor (L.) Moench". https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:422090-1. 
  14. ↑ "Perennial Sorghum". The Land Institute. https://landinstitute.org/our-work/perennial-crops/perennial-sorghum/. 
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  16. ↑ "sorghum (n.)". Online Etymology Dictionary. https://www.etymonline.com/word/sorghum. 
  17. ↑ "Definition of Milo". https://www.merriam-webster.com/dictionary/milo. 
  18. ↑ "Definition of Durra". https://www.merriam-webster.com/dictionary/durra. 
  19. ↑ Nicholson, Zoe (8 April 2018). "A Sweet Security: White African Sorghum". https://www.thecarolinagoldricefoundation.org/news/2018/4/18/a-sweet-security-white-african-sorghum. ⇐
  20. ↑ Basinger, Ryan (31 May 2017). "Grain Sorghum for Deer". https://deerassociation.com/grain-sorghum-deer/. 
  21. ↑ "Sorghum". https://oklahoma.agclassroom.org/resources/agricultural-facts/ag-facts/sorghum/. 
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  32. ↑ "Sweet Sorghum". Sweet Sorghum Ethanol Producers. http://sseassociation.org/SweetSorghum/default.aspx. 
  33. ↑ Hyde, James F.C. (1857). The Chinese Sugar-Cane: Its History, Mode of Culture, Manufacture of the Sugar, Etc. with Reports of Its Success in Different Portions of the United States, and Letters from Distinguished Men. Boston: J. P. Jewett. https://books.google.com/books?id=fA_tOyZnXyUC&q=sorghum+sugar+syrup+china. 
  34. ↑ Hyde, James F.C. (1857). The Chinese Sugar-Cane: Its History, Mode of Culture, Manufacture of the Sugar, Etc. with Reports of Its Success in Different Portions of the United States, and Letters from Distinguished Men. Boston: J. P. Jewett. p. 11. https://books.google.com/books?id=fA_tOyZnXyUC&q=sorghum+sugar+syrup+china. 
  35. ↑ Mulhollem, Jeff (10 August 2020). "Flavonoids' presence in sorghum roots may lead to frost-resistant crop". Pennsylvania State University. https://news.psu.edu/story/627935/2020/08/10/research/flavonoids-presence-sorghum-roots-may-lead-frost-resistant-crop. "sorghum is a crop that can respond to climate change because of its high water- and nitrogen-use efficiency" 
  36. ↑ Danovich, Tove (15 December 2015). "Move over, quinoa: sorghum is the new 'wonder grain'". The Guardian. https://www.theguardian.com/lifeandstyle/2015/dec/15/sorghum-wonder-grain-american-food-quinoa. 
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  39. ↑ "Tapping into Sorghum's Weed Fighting Capabilities to Give Growers More Options". https://www.ars.usda.gov/news-events/news/research-news/2010/tapping-into-sorghums-weed-fighting-capabilities-to-give-growers-more-options/. 
  40. ↑ "Sorghum – Section 4: Plant Growth and Physiology". https://grdc.com.au/resources-and-publications/grownotes/crop-agronomy/sorghumgrownotes/GrowNote-Sorghum-North-04-Physiology.pdf. 
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  49. ↑ Guo, Chunshan; Cui, Wei; Feng, Xue; Zhao, Jianzhou; Lu, Guihua (2011). "Sorghum insect problems and management". Journal of Integrative Plant Biology 53 (3): 178–192. doi:10.1111/J.1744-7909.2010.01019.X. PMID 21205185. 
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  53. ↑ Bandyopadhyay, Ranajit; Frederickson, Debra E.; McLaren, Neal W.; Odvody, Gary N.; Ryley, Malcolm J. (April 1998). "Ergot: A New Disease Threat to Sorghum in the Americas and Australia". Plant Disease 82 (4): 356–367. doi:10.1094/PDIS.1998.82.4.356. PMID 30856881. Bibcode: 1998PlDis..82..356B. 
  54. ↑ Waniska, R. D.; Venkatesha, R. T.; Chandrashekar, A.; Krishnaveni, S.; Bejosano, F. P.; Jeoung, J.; Jayaraj, J.; Muthukrishnan, S. et al. (1 October 2001). "Antifungal Proteins and Other Mechanisms in the Control of Sorghum Stalk Rot and Grain Mold". Journal of Agricultural and Food Chemistry 49 (10): 4732–4742. doi:10.1021/jf010007f. PMID 11600015. Bibcode: 2001JAFC...49.4732W. 
  55. ↑ Paterson, Andrew H.; John E. Bowers; Remy Bruggmann; Inna Dubchak; Jane Grimwood; Heidrun Gundlach et al. (2009-01-29). "The Sorghum bicolor genome and the diversification of grasses". Nature 457 (7229): 551–556. doi:10.1038/nature07723. PMID 19189423. Bibcode: 2009Natur.457..551P. 
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  94. ↑ Klaniczay, Gábor (1990). The Uses of Supernatural Power: The Transformation of Popular Religion in Medieval and Early-Modern Europe. Princeton: Princeton University Press. pp. 129–130. ISBN 978-0-6910-7377-4. 

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